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An international journal to expedite plant systematic, ptiytogeographical 
and ecological publication 

Vol. 81 August 1996 No. 2 


HIRSHBERG. J. & D. CLEMONS, Vascular plants of ihe Cuyamaca 

Mountains, California 69 

TURNER, B.L., Perynieniiini basaseacliicaniitn (Asteraceae, Hclianlheae), a 

new species from Chihuahua, Mexico 103 

TERRELL, E.E., Taxonomic notes on Texan and Mexican species of Hedyotis 

and Houslonia (Rubiaceae) 108 

TURNER, B.L., Vernonia hoquerona (Asteraceae), a new species from 

Chiapas, Mdxico 115 

TURNER, B.]^.,Ageratiiui bobjansenii (Asteraceae: Eupatoricac), a new species 

from Durango, Mexico 118 

LAFERRIERE, J.E., Prionosciadiiun saraviki (Apiaccae): A new species from 

the Sierra Madre Occidental, Mexico 121 


of culm anatomical characters in the species of Bouieloitu Liigasca (Poaccae: 

Eragroslidcae) ; 1 24 

REVEAL, J.L., Aridae, a new subclass of Liliopsida 142 

Phytologia Memoirs 1 1 available 144 


MAY 2 6 1997 


PHYTOLOGIA (ISSN 00319430) is published monthly with two volumes per year by 
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Phytologia (August 1996) 81(2)69-102 



Jerilyn Hirshberg 

JBH Biological Surveys, P.O. Box 2, Julian, California 92036 U.S.A. 


Duffie demons 

c/o Botany Department, San Diego Natural History Museum, P.O. Box 1390, San 
Diego, California 92112 U.S.A. 


A total of 996 taxa are included in this checklist of the vascular plants of 
the Cuyamaca and Laguna mountains of San Diego County, California. The 
introduction includes information on climate, geology, and patterns of 
biogeographical disjunction in this area. 

KEY WORDS: vascular plants, Cuyamaca and Laguna mountains, California 

This checklist is based on Plants of Montane San Diego County compiled by 
Duffie Clemons (1986) for the California Nati\e Plant Society and the San Diego 
Natural History Museum. The area covered here includes the Cuyamaca and Laguna 
mountains of central San Diego County, California and their adjacent foothills. 
Altitudes range from approximately 910 meters (3,000 feet) on the eastern and western 
boundaries to a high of 1970 meters (6,500 feet) on Cuyamaca Peak. 

The Cuyamacas and Lagunas are pardilel, north-south trending ranges that 
con\erge in the vicinity of Cuyamaca Lake. Corte Madera Mountain and Guatay 
Mountain, which are kxxited slightly south of these two ranges, Volcan Mountain, 
located immediately north, and Poser and Viejas mountains and Mesa Grande, which 
lie slightly west of the Cuyamacas, are also included. The small settlements of 
Buckman Springs and San Felipe lie on the eastern boundary. Banner lies 
approximately five kilometers (three miles) within this boundary. The distance from 
the western to the eastern boundary is approximately 24 kilometers (fifteen miles), and 
40 kilometers (25 miles) from the northern to the southern boundaries. 


70 PHYTOLOGIA August 1996 volume 81(2):69- 102 


The Cuyamaca Mountains receive more rain than any other area in southern 
California of comparable elevation, with the exception of Palomar Mountain. The high 
rainfall in these mountains is apparently due to their location, within forty miles of the 
coast and parallel to it, with no intervening ranges to catch the rainfall (Beauchamp 
1972). Average annual rainfall totals cited by Wiggins ( 1929) at vanous elevations in 
the mountains of San Diego County, based on penods ranging from 2 to 69 years 
include the following: 


100.93 cm 

40.37 in 


59.94 cm 

23.60 in 

Volcan Mt. 

88.25 cm 

35.30 in 

San Felipe 

59. 18 cm 

23.30 in 


73.71 cm 

29.02 in 

Mt. Laguna 

57.58 cm 

22.67 in 

Additional data from the County of San Diego Department of Flood Control 
indicates that yearly rainfall averages approximately 1 14.3 centimeters (45 inches) on 
Mount Palomar and 77.47 centimeters (30.5 inches) at Mesa Grande (Allan Rand, 
County of San Diego climatologist, pers. comm.). The mountains of San Diego 
County receive a significant portion of their rainfall during the summer months with 
Hot Springs Mountain and the Laguna Mountains receiving the most summer rain 
(Raven & Axelrod 1978). Summer rain may help to offset low winter rainfall during 
drought years, and it is also possible that a more regular pattern of rainlall throughout 
the year may affect vegetation in other ways. Snow and cold temperatures occur in 
San Diego County mountains in winter. The highest average annual snowfall of 
approximately 203 centimeters (80 inches) occurs on Mount Laguna (Lower 1978). 

The Cuyamaca and Laguna mountains are part of the Peninsular Ranjge which 
extends from Mount San Jacinto in Riverside County almost to the southern tip of Baja 
California. The Peninsular Range is underlaid by the intrusive igneous geologic 
formation known as the Southern California Batholith. The lighter phases of plutonic 
(intrusive igneous) rock which contain a higher proportion of light-colored minerals 
are called granitic, and the darker rocks which contain more dark-colored minerals are 
known as gabbro. 

The Cuyamaca Mountains, Poser, Viejas, and Gualay mountains. Mesa Grande, 
portions of Corte Madera Mountains and Garnet Peak in the Laguna Mountains are 
underlaid by gabbro. Chiquilo Peak, Stonewall Peak, p<.)rtions of Corte Madera 
Mountain, and part of the Volcan Range are granitic. Micaceous schist, a melamorphic 
rock, predominates on the eastern escarpment of the Laguna and Volcan mountains. 

Soils denved from extremely light granitic rtK-k arc often sandy and sterile. Soils 
derived from extremely dark gabbroic rock often form heavy clays that contain highly 
charges ions of magnesium and iron. Clay soils have a high water-holding capacity, 
and are also p(^x)rly aerated during the winter months. Although both of these extreme 
soil types are unfavorable for many common species, other less common species are 
able to tolerate Ihem. from competition from the more dominant species may 
also be a factor which enables some rare plants lo persist on these soil tvpes 
(Oberbaucr 1993). 

Hirshberg & demons: Cuyamaca and Laguna Mountains Oora 71 

Plant communities uithin the study area range from coastal sage scrub and 
chaparral on the western boundary to mixed coniferous forest at the higher elevations, 
and pinyon-juniper utxxlland near the eastern boundary. Extensive mountjun 
meadows that retain standing water until late spnng are present at higher elevations in 
the Laguna Mountains. Cuyamaca Lake was called La Laguna Que Se Seca (the lake 
that dries up) by the early Spanish settlers. The valley in which this lake is located 
extends from the cismontane to the desert slope of the range, a distance of 
approximately eight kilometers (five miles) in this area. A dam constructed in 1886 at 
the western end of the valley now maintains a more permanent lake currently used as a 

Patterns of Biogeographical Disjunction 

A total of 996 taxa are included in this checklist. More than fifty plant species 
regarded as rare by the California Nati\ e Plant Society (Skinner & Pavlik 1994) occur 
in the Cuyamaca and Laguna mountains. Staking cases of disjunct distribution which 
appear to be relictual also occur in the mountains of San Diego County. These are 
species which are common in northern California, but which are known in southern 
California exclusively, or almost exclusively from this area. It is probable that these 
species were more widely distributed in southern California dunng cooler, wetter 
climatic regimes, and have been able to persist here because of "special soil and 
precipitation factors" (Munz 1974). Some disjuncts which have evolved into 
subspecies or varieties endemic to this region are regarded as rare taxa in their own 

Elisabeth Crow (later Norland), a student of Dr. P.A. Munz at Pomona College, 
noted in her Master's thesis (1931) that high rainfall and gentle topography are 
probably among the factors v\ hich have allowed many northern species to survive here 
after being extirpated from the intervening ranges. Other climatic factors may include 
summer rain and snow in winter. Crow (1931) also noted that di.sjuncts occur in 
cismontane and coastal as well as montane areas of both San Diego and Riverside 
counties where "extreme moist conditions pjersist". E.xamples which she cited include 
Rhododendron occidenlale (Torrey & A. Gray) A. Gray, Rosa gyninocarpa, 
Ceanotlius foliosiis C. Parrv', and Ophioglossiim californicuni. 

Other observ ations also suggest that substrates where "extreme moist conditions 
persist" play a role in disjunction. Many of the disjunct taxa which cx;cur in the 
Cuyamaca Mountains are restricted to gabbro soils. Polygonum parryi E. Greene, 
known in southern California only from small populations at Cuyamaca Lake and on 
Cuyamaca Peak, txjcurs on vernally wet clay soils. Epilobiiim minutiitn Lehm., 
Aspidolis densa (Brackencr) Lcllingcr, and Moeliringia nuicrophylla are also gabbro- 
restricted, and the latter two are as.stx-iatcd u iih serpentine soils in northern California. 
Like gabbro, serpentine weathers to form soils which have a high water-holding 
capacity, and are also unfavorable for many plant species (Raven & Axclrcxi 1978). 

Two additional disjuncls which arc known in southern California onl\ from the 
Cuyamaca Mountains arc ass(x;iatcd with vernal p<xils and wet meadows near 
Cuyamaca Lake: Navcirrelia higelina E. Greene and Downingia concolor E. Greene 

72 PHYTOLOGIA August 1996 volume 81(2):69- 102 

subsp. brevior McVaugh. Several other disjuncls which are more widely distributed 
in both the Cuyamaca and Laguna mountains arc also associated with vernal pools and 
dr>'ing meadows which may, or may not be, underlaid by gabbro. These include 
Mimiartia piisilla (S. Watson) Mattf., Thermopsis macrophylla Hook. & Am. var. 
seniola Jepson, Delphittiiim hesperium A. Gray subsp. cnyaniacae (Abrams) Harlan 
Lewis & Epling, and Limnanthes gracilis Howell subsp. parishii (Jepson) 

In the Cuyamaca foothills, Mimulus congdonii Robinson, which is disjunct from 
the Sierra Nevada and Ventura County, occurs on damp, sandy, granitic soils near 
Descanso. Ceanothus foliosus var. foliosus, which occurs on gabbro soils in 
chaparral in both foothill and montane areas of the Cuyamacas, is also a disjunct which 
is more widely distnbuted in northern California. FiUigo arizonica A. Gray and 
Harpagoiiella paUneri k. Gray are coastal -desert disjuncts which are restricted to clay 
soils on the cismontane slope, but not necessanly so in desert areas. 

In general, distribution patterns suggest that many disjuncts have compensated for 
gradually decreasing rainfall by retreating to habitats and substrates which are moist, 
and also unfavorable for more dominant plant species. A more complete list of 
disjuncts occurring in this area, a life form spectrum, and the number of non-native 
species and genera per family are included in appendices at the end of this checklist. 


All taxa included in this checklist are referable to the San Diego County synoptic 
collection housed at the herbarium of the San Diego Natural History Museum (SD). 
Locations in the Cuyamaca and Laguna mountains are also documented in the main 
collection at SD and, in some instances, in herbana housed at the University of 
California, Riverside (UCR) and at Rancho Santa Ana Botanic Garden, Claremont 

New locations, not cited by Beauchamp (1986), are included for a few taxa. 
County range extensions are documented with collector's name and number, and 
herbarium accession number in the ChecklisI of the Vascular Plants of San Diego 
County (Simpson, et al. 1994). Scientific nomenclature follows Hickman (1993). 
Common names are taken from Beauchamp ( 1986) and Hickman ( 1993). 

Vascular Plants of the Cuyamaca and Laguna Mountains 

Symbols used in this list are explained at the end of the list. 


ASPLENIACEAeSplccnwort Family 

Asplenium vesperliniini Maxon. Corlc Madera Ranch, Cedar Creek 

Hirshberg & demons: Cuyamaca and Laguna Mountains llora 73 

AZOLLACEAE/Mosquito Fern Family 
Azolla filiculoides Lam. Pacific mosquito fern 

BLECHNACEAE/Deer Fern Family 
Woodwardia fimbriala Smith Giant chain fern 


Pteridium aquilinum (L.) Kuhn var. piibescens (L.) Underw. 

Cystopleris fragilis (L.) Bemh. Bntlle bladderfern 
Dryopteris arguta (Kaulf.) Max on Coastal wood fem 

Polystichiim imbricmis (D. Eaton) D.H. Wagner subsp. curtum (Ewan) D.H. Wagner 
Canyon sword fem 

EQUISETACEAE/Horsetail Family 

Equisetum arveiise L. Common horsetail 

Equisetum hymale L. subsp. affine (Engelm.) Calder & R.L. Taylor 

Equisetum laevigatum A. Braun 

Equisetum telematela Ehrh. subsp. braunii (Milde) R.L. Hauke 

ISOETACEAEyQuillwort Family 
Isoetes fwwellii Engelm. Corte Madera Ranch 
Isoetes nuttallii Engelm. Cuyamaca Peak 
Isoetes orcuttii A. A. Eaton Corte Madera Ranch 

MARSILEACEAE/Marsilea Family 

Marsilea vestita Hook. & Grev. subsp. vestita Hairy cloverfem 

Pilularia americana A. Braun Amencan pillwort 

POLYPODIACEAE/Polypody Family 

Polypodium califoniicum Kaulf. California polypody 

Adianlumjordanii C. Mueller Califomia maidenhair 
Aspidolis califoniica (Hcxik.) Copcl. California lacefem 
Aspidolis deiisa (Brackcncr) Lcllingcr Cuyamaca Peak; Lacefem 
Cheilanthes develandii D. Eaton Clc\cland's lipfem 
Clieilanlhes covillei Mdxon Coville's lipfern 
Pel Idea aiidroniedifolid (Kaulf.) Fee Coffee fem 
Pellaea nmcrunala (D. EEaton) D. Elaton \ar. mucronata 

Penlograinimi triangularis (KduW.) G. YaLskicvych, Windham, & Wollenweber subsp. 
triangularis Goldenback fern 

Selaginella asprella Maxon Bluish mosslern 
Selaginella bigelovii L. Undcru. Bigclou's mossfcm 

74 PHYTOLOGIA August 1996 volume 81(2):69- 102 


CUPRESSACEAE/Cyprcss Family 

'Calocedrus decurretis (Torrey) Flonn Incense cedar 

'Cupressus arizonica E. Greene subsp. arizonlca [C. stephensonii C. WolH 

Cuyamaca cypress 
'Cupressus forbesii Jepson Tecate cypress 
'Juniperus californica Carriere California juniper 

EPHEDRA CEAE/Ephedra Family 

'Ephedra californica S. Watson thing Valley; Desert tea 

PINACEAE/Pine Family 

^Abies coticolor (Gordon & Glend.) Lindley White fir 

^Piniis coulteri D. Don Coulter pine 

'Pinus jeffreyi Grev. & Balf. Jeffrey pine 

'Pinus lamberliatui EXiuglas Sugar pine 

^Piniis monophylla Torre> & Fremont Chariot and Indian canyons 

'Pinus ponderosa Laws. Ponderosa pme 

'Pinus quadri/olia Pari. 

'Pseudotsuga macrocarpa ( Vasey) Mayr Big-cone Douglas fir 



ALISMATACEAE/Water-pIantam Family 
^Alisnia planlago-aqualica L. Julian and Wynola 

CYPERACEAE/Sedge Family 

^Carex alma L. Bailey Sturdy sedge 

^Carex globosa Boolt 

^Carex mullicaulis L. Bailey 

^Carex praegracilis W. Boott 

^Carex sartwelliana Olney Cuyamaca Peak 

^Carex senla Boott 

^Carex spissa L. Bailey 

^Carex subfusca W. B(X)lt 

'Cypenis parisliii Britton Parish's umbrella-sedge 

'Cyperus squarrosus L. 

^Eleocluiris acicularis (L.) Rcx;mcr & Schultes \ar. bella Piper 

^Eleocharis nmcrostachya B niton 

^Eleocharis inonlevideiisis Kunlh 

^Eleoctiaris parisliii Bntlon 

^Eleocharis paucijlora (Light.) Lmk 

^Scirpus aculiis Bigelow \ar. occidentalis (S. Watson) Beetle 

^Scirpns ainericanus Pcrs. Amcncan bulrush 

''Scirpus niicrocarpus C. Presl Small-lruit bulrush 

Hirshberg & Clemons: Cuyamaca and Laguna Mountains Hora 75 


Vm missouriensis Null. Rcx;ky Mountain blue flag 

^SisyrinchiumbellumS. Watson Biue-eyed-grass 

JUNCACEAE/Rush Family 

*Juncus ambiguus Guss. 

*'Juncus balticus Willd. Baltic rush 

'Juncus bryoides F.J. Hcrm. One-llower dwarf rush 

^ J uncus bufonius L. var. bufonius Toad- rush 

*Juncus bufonius L. var. occidenlalis F.J. Herm. 

^Juncus dubius Engelm. Manposa rush 

^Juncus ejfusus L. var. pacificus Fern. & Wieg. 

V uncus luciensis B. Ertter 

^'J uncus macrophyllus Gov. 

^'Juncus mexicanus Willd. 

^ J uncus oxyineris Engelm. 

*'Juncus patens E. Meyer 

^Juncus pluieocephalus Engelm. var. paniculatus Engelm. 

^Juncus rugulosus Engelm. 

^Juncus tenuis Willd. 

* J uncus tiehmii B. Ertter 

^ J uncus torreyi Gov. 

^Juncus xiphioides E. Meyer 

^Luzula comosa E. Meyer Cuyamaca Peak; Hairy wood-rush 

JUNGAGINAGEAE/Arrowgrass Family 

'Lilaea scilloides (Poiret) Hauman Flowering quillwort 

LEMNAGEAE/Duckweed Family 
*'Lemna minor L. Lesser duckw eed 
^Lemna minitscula Herter Least duckweed 

LlUAGEAEyLily Family 

^Allium ainplectens Torrey Paper onion 

^Allium campanulatum S. Watson Sierra onion 

^Allium Jimbriotu/n S. Watson var. Jlmbrialuni Desert onion 

^Allium luieniatocliiton S. Watson Red-skin onion 

^Allium parryi S. Watson Guyamaca Lake, Filaree Rat 

^Allium peninsulare Lemmon \ ar. peninsulare Red-flower onion 

"Asphodelus Jistulosus' L. Descanso; Hollow -stem asphodel 

*Bloonieria crocea (Torrey) Gov . Gommon goldenstar 

*Brodiaea orcullii (E. Greene) Baker Orcutt's brodiaea 

Hirodiaea terrestris Kellogg subsp. kernensis {Hoo\c\) T. Niehaus 

^Calochortus albus Benth. Fairv lantern 

*Coloc/u)rlus concolor (Baker) Purdy Golden-bowl manposa lily 

*Caloclu>rtus dunnii Purdy Dunn's manposa lily 

^Calochorliis inveniisliis E. Greene 

*Colochonus splendens Benth. Lilac manposa lily 

K^dlochortiis weedii A\ph. Wcxvl \ar. weedii 

K^hlorogalinn parvijhmim S.^n Poser and Vicjas inouniains 

^Dichelostenniia capilaliiin Alph. W(hk1 subsp. cdpiUiJiiin 

76 PHYTOLOGIA August 1996 volume 81(2):69- 102 

*Fritillaria bijlora Lindley var. bijlora Viejas Mountain 

'Liliu/n humboldlii Roezl & Leichtlin subsp. ocellatum (Kellogg) Thome Ocellated 

tiger lily 
'Lilium pardalinum Kellogg subsp. pardalinum Leopard lily 
'Muilla tmrilima (Torrey) S. Watson Poser Mountain 
*'Nolina cistnonuma Dice. Viejas Mountain 
^Nolina parryi S. Watson Parry's bear-grass 
*'SmiUicina stellata (L.) Desf. False Solomon's seal 
'Yucca schidigera K.E. Ortgies Spanish dagger 
^Yucca Whipple ilovxty Whipple's yucca 
'Zigadenus Jreniontii (Torrey) S. Watson Viejas Mountain 
'Zigadenits venenosus S. Watson var. venenosus Death camas 

ORCHIDACEAE/Orchid Family 

^Corallorhiza maculata Raf. Spotted coralroot 

^Epipactis gigatitea Hook. Giant stream orchid 

^Piperia elongala Rydb. Cuyamaca RSP 

^Piperia leptopelala Rydb. Cuyamaca RSP 

^Piperia transversa Suksd. Cuyamaca RSP 

^Piperia utuilascensis (Sprengel) Rydb. Cuyamaca RSP 

^Platantfiera leucostachys Lindley Cuyamaca RSP 

*'Spiranthes porrifolia Lindley Ladies' tresses 


^Achtiatherwn coronaliun (Thurber) Barkworth Giant needlegrass 

^Achnatherum fiyme noide s {Roemer & Schultes) Barkworth 

^Achnatfierum lemmonii (Vasey) Barkworth Lemmon's needlegrass 

^Achnalherutn occidentalis (Thurber) Barkworth subsp. californicum (Merr. & Burtt 

Davy) Barkworth Cuyamaca RSP 
^Achnatherum occidentalis (Thurber) Barkworth subsp. occidentalis Laguna 

^Achnatheniin speciosum (Tnn. & Rupr.) Barkworth 
^Agrostis exarata Trin. Spike bcntgrass 
^Agrostis pallens Tnn. Leafy bcntgrass 
*'Agrostis scabra Willd. Rough benlgrass 
^Agrostis stolonifera' L. Creeping bcntgrass 
^Agrostis viridis Gouan Water bcntgrass 
'Aira caryophyllea' L. Silver hairgrass 
^Alopeciiriis aequalis Sobol. Short-awn fo.xtail 
'Aloperurus carolinianus Walter Carolina foxtail 
'Alopeciirns saccatiis Vasey Pacific foxtail 
*Aristida adsceiisionis L. Vicjas Mountain 
^'Aristida purpurea Null. var. longisela (Slcudcl) Vasey 
^Aristida purpurea Nult. var. nealleyi (Vasey) K.W. AUred 
^Arislida purpurea Null. var. purpurea Vicjas Mountain 
*Avena barbala' Link Slender u ild oat 
^Avenafatua' L. 

^Avena saliva' L. Culti\ alcd oat 

^Holhriochloa barbiiuxtis (Ux^.) Hcrlcr Viejas Mountain 
'Bouleloiia barbala Lag. \ ar. barbala Six- weeks grama 
^liouleloua curlipendula ( Michaux) Torrey Side-oats grama 

Hirshberg & Clcmons: Cuyamaca and Laguna Mounlains flora 77 

'Briza minor L. Corte Madera Ranch; Little quaking-grass 

^lironuLS arenariiis Labill. Australian chess 

^Bromus carinaliis Hook. & Am. var. cariiuitiis California brt)me 

'Bromus diandriLs' Roth Ripgut brome 

^Bromus grandis (Shear) A. Hitchc. Grand brome 

'Bromus hordeaceus' L. Smooth brome 

'Bromus japonicus' Murr. Japanese brome 

'Brotnus madriteiisis L. subsp. riihens (L.) Husnot Red brome 

^Bromus orcuttianus (Shear) A. Hitchc. Orcutl's brome 

'Bromus tectorwn L. Cheat-grass brome 

'Bromus trinii Desv. Chilean chess 

^CaUimagrostis koelerioides Vasey Dense reedgrass 

^Crypsis vaginiflora (Forsskal) Opiz 

^Cynodon dactylon (L.) Pers. Bermuda-grass 

'Cynosunis echinalus' L. Santa Isabel, Stonewall Peak 

^Dactylis glomerala L. Orchardgrass 

^DaiiOionia caUfornica Bolander var. americaiia (Scnbner) A. Hitchc. California 

^Dantlionia unispicata (Thurber) Macoun One-spike oatgrass 
'Descfiampsia danthonioides (Trin.) Benth. Hairgrass 
^Deschainpsia elongata (Hook.) Benth. Slender hairgrass 
^Distichlis spicala (L.) E. Greene Salt-grass 

^Elymus elynioides (Raf.) Swezey subsp. brevifoHus (J.G. Smith) Barkworth 
^Elymus glaiiciis Buckley subsp. glaucus Blue rye 
^Elymus glaucus Buckley suhsp. jepsonii (Burtt Davy) Gould 
^Elymus glaucus Buckley subsp. virescens (Piper) Gould 
^Elymus multisetus (J.G. Smith) Burtt Davy 
*'Elymus tracliycaulus (Link) Shinncrs subsp. Irachycaulus 
^Elylrigia intermedia (Host) Nevski subsp. intermedia' 
^Elytrigia pontica (Podp.) Holub. subsp. poH//ra* 
^Eragrostis curvula (Shrader) Nees var. conferta' Nees Descanso 
^Eragrostis minor Host 
^Festuca occidentalis Hook. Western fescue 
^Eestuca rubra L. Red fuscue 

'Gastridium ventricosum' (Gouan) Schinz & Thell. Nitgrass 
^Holcus lanatiLs' L. Creeping vehct-grass 
^Hordeum brae hy ant he rum Ne\ ski subsp. brachyantlientm 
^Hordeum brachvanthenan Ne\ski subsp. californicum (Co\as & Stebb.) V. 

Bothmer, N. Jacobscn, & O. Scberg 
'Hordeumjubatum L. 

'Hordeum nuirinum Hudson subsp. gussoneanum' (Pari.) Thell. 
^Hordeum murinum L. subsp. glaiiciim' (Steudcl) Tzvele\ 
'Hordeum murinum L. subsp. leporiniim' (Link) Arcang. 
'Hordeum vulgare' L. Culti\atcd barley 
^Koeleria macranl/ia (Ledeb.) J. A. Schultcs Junegrass 
'Uunarckia aurea' (L.) Mocnch Golden -top 
^Levmus x wulliflorus (Gould) Barkworth & D.R. Dewey 
^lx'\inus iriiicoides (Buckley) Pilger Beardless rye 
^Lolium perenne' L. Perennial ryegrass 
'Lolium temulenlum' L. Milk Ranch Road (UCR) 
*'A/f//'<Y;/r///e'.vct'm' Scnbner Tall mclic 

78 PHYTOLOGIA August 1996 volume 81 (2):69- 102 

^Melica imperfecUiTun. 

^Muhlenbergia asperifolia (Nccs & Meyen) L. Parodi 

^Muhlenbergia microspenm (DC.) Kunth Little muhiy 

^Muhlenbergia richardsonis (Tnn.) Rydb. Mat muhiy 

^Muhlenhergia rigens (Benth.) A. Hitchc. Deergrass 

^Nasella cernua (Stebb. & Love) Barkworth East Mesa 

*'Nasella lepida (A. Hitchc.) Barkworth Foothill needlegrass 

^Nasella pulchra (A. Hitchc.) Barkworth Purple needlegrass 

^Panicum acuminatum Sw. var. acuminatum Pacific witchgrass 

*Panicum capillare L. Common witchgrass 

^Paspalum distichum L. Common knotgrass 

^Pennisetum setaceum' Forsskal 

^Pfmlaris aquatica' L. Harding grass 

^P/mlaris arundinacea' L. Big Laguna Lake; Reed canary grass 

'Pfmlaris minor Retz Little-seed canary grass 

^Phleutn prateme' L. Timothy 

^Piptathenun miliaceum' (L.) Cosson Smilo 

'Poa annua' L. Annual bluegrass 

^Poa atropurpurea Scribner Laguna and Corte Madera mountains 

^Poa bulbosa' L. Bulbous bluegrass 

^Poa compressa' L. Canada bluegrass 

*Poa fiowellii Vasey & Scnbner Cuyamaca RSP 

^Poa pratensisL. subsp. prateiusis' Kentucky bluegrass 

^'Poa secunda J.S. PresI subsp. secunda Malpais bluegrass 

'Polypogon interrupt i4s' Kunth Ditch beardgrass 

'Polypogon monspeliensis' (L.) Desf. Rabbi t-ftx)t grass 

^Schismus barbatus' (L.) Thcll. 

'Secale cereale L. Descanso; Common rye 

^Sorghum halepense' (L.) Pers. Johnson grass 

^Splienopliolis obtusata (Michaux) Scribner Cuyamaca Mountains 

^Sporobolus airoides (Torrey) Torrey Corte Madera Ranch 

^Sporobolus cryptandrus (Torrey) A. Gray Sand dropsced 

'Triticum aestivum' L. Laguna Lakes; Cereal wheat 

^Vulpia bromoides' (L.) S.F. Gray 

Wulpia microstachys (Nutt.) Benth. var. cUiata (Beal) Lonard & Gould 

'Vulpia microstachys (Nutt.) Benth. var. confusa (Piper) Lonard & Gould 

^Vulpia microstachys (Nutt.) Benth. var. pauciflora (Beal) Lonard & Gould 

^Vulpia myuros{L.) C. Gmclin var. hirsuta' (Hackell) Asch. & Graebner 

""Vulpia ocloflora (Walter) Rydb. var. hirlella (Piper) Henrad 

'Vulpia octoflora (Walter) Rydb. var. ocloflora 

^Polamogeton foliosus Raf. \m. Joliosus Lealy pondwecd 
^Potamogeton illinoensis Morong Shining pondwecd 
^Potamogeton nodosus Poirct Long-leal pondwecd 
^Potamogeton pectinatus L. Fcnncl-lcal' pondwecd 
^Potamogeloii pusilltts L. var. piisilliis Small pondweed 

TYPHACEAE/Cal-tail Family 

^Typha domini;ensis Pcrs. Tulc cat- tail 

^typtia lalifol'ia L. Soft Hag 

Hirshbcrg & Clcmons: Cuyamaca and Laguna Mountains (lora 79 

ZANNICHELLIACEAE/ Horned pondweed Family 
^Zannicliellia paliLslris L. Horned pondweed 


ACERACEAE/Maple Family 

^Acer nmcrophylhun F*ursh Big-leal maple 

AIZOACEAE/Carpetweed Family 
'Cypselea humifusa Turpin 

AlVLARANTHACEAE/Amaranth Family 
*A/naranllius albiis L. While tumbleweed 
^Aniaranthus bliloides S. Walson Prostrate amaranth 
^Aniaranlhus californiciis (Moq.) S. Watson 
*Atnaranlfii4s relrojlexiis' L. 


^Malosnui laurina (Nutt.) Abrams Laurel sumac 

^Rhus ovata S. Watson Sugar bush 

'Rhus trilobata Torrey & A. Gray Basket bush 

"Toxicodendron diversilobiun (Torrey & A. Gray) E. Greene 


^Angelica lomentosa S. Watson 

"Anlhrisciis caucatis' M. Bieb. Cuyamaca Lake 

'Apiastrimi angustijoliuni Nutt. Mock parsley 

"Apiuin graveolens L. Wild celery 

^Berula erecta (Hudson) Gov. Cut-leal' water parsnip 

"Bowlesia incana Ruiz Lopez & Pa\ on Corie Madera Ranch 

^Ciciita douglasii (DC.) J. Coulter & Rose Water-hemlock 

"Coniion nuiculatiun L. Mount Laguna; Poison hemkx^k 

^Daucus carola L. Wild carrot 

'Daucus pusillus Michaux Rattlesnake weed 

^Hydrocotyle ranunculoides' L.f. Marsh pennywort 

^Lotmitiiim dasycarpum (Torrey & A. Gray) J. Coulter & Rose subsp. dasycarpum 

Bladder parsnip 
^Lomaliinn lucid iitn (Torrey & A. Gray) Jepson Shiny lomatium 
^Oenonlhe sarweiitosa J.S. PresI Pacific oenanlhe 
^Osinorhiza Imicliypoda Torrey California sweet-cicely 
^Osinorhizci chilensis Hcx>k. & Arn. Mountain sweet-cicely 
^Faslinaca saliva L. Common parsnip 
^Perideridia parishii (J. Coulter & Rose) Nelson & J.F. Macbr. subsp. lalifolia (A. 

Gray) Chuang & Constance 
*'Sanicula arguta J. Coulter & Rose Sharp-tcx)th sanicle 
^Saniciila bipinnatijida Hcx^k. Puiplc sanicle 
^Soniciila crassicaulis DC. Pacific sanicle 
\Sanicula graveolens DC. Sierra sanicle 
^Saniciila iiiberosa Toney Turkc\-pca sanicle 

80 PHYTOLOGIA August 1996 volume 81 (2):69- 102 

*Tauschia arguta (Torrey & A. Gray) J.F. Macbr. 
^Taiischia parishii (J. Coulter & Rose) J.F. Macbr. 
'Torilis arvetisis' (Hudson) Link. Cuyamaca RSP 
'Yabea microcarpa (Hook. & Am.) Koso-Polj. 

APOCYNACEAE/Dogbane Family 
"Apocynum androsaeinifolium L. 
^Apocynum cannabinum L. Indian hemp 

*'Asclepias cali/ornica E. Greene Woolly milkweed 
*'Asclepias eriocarpa Benth. Indian milkweed 
^Asclepias erosa Torrey Desert milkweed 
^Asclepias fascicularis Decne. Narrow-leaf milkweed 

^Achillea filipetidulina Lam. Pine Hills; Fem-leaf yarrow 
*" Achillea millefolium L. Yarrow 
^Acourtia microcepluila DC. Scapellote 
^'Agoseris grandiflora (Nult.) E. Greene 
*Agoserisheterophylla (Nutt.) E. Greene 
^Agoseris retrorsa (Benth.) E. Greene 
'Ambrosia acanthicarpa Hook. Annual bur-weed 
'Ambrosia dumosa (A. Gray) Payne White bur-sage 
^Ambrosia psilostachya DC. Western ragweed 
^Atiisocoma acaulis Torrey & A. Gray Scale-bud 
'Anthemis colula' L. Mayweed 
'Artemisia cali/ornica Less. Coastal sagebrush 
^'Artemisia douglasiana Besser Douglas mugwort 
^Artemisia dracunculiis L. Tarragon 
'Artemisia ludoviciana Nutt. subsp. albula (Wooton) Keck 
'Artemisia ludoviciami Nutt. subsp. ludoviciana Common mugwort 
'Artemisia tridentata Nutt. subsp. tridenlala 
^'Asler bernardinus H.M. Hall San Bernardino aster 
^Aister chilensis Nees Chile aster 

'Daccluiris salicifolia (Ruiz Lopez & Pavon) Pers. Mule-fat 
'Baccluiris sarothroides A. Gray Descanso; Broom baccharis 
'Baccharis sergiloides A. Gray Squaw waterueed 
*Bidens pilosa L. var. pilosa' Beggar's tick 
^Blennosperma nanum (Hcxik.) S.F. Blake var. nanum 
'Brickellia cali/ornica^ (Torrey & A. Gray) A. Gray 
'Centaurea nuuidosa'^Lnm. Julian; SpcMtcd knapweed 
'Centaurea melilensis' L. Tocaiolc 
'Centaurea solslilialis' L. Yellow star-lhisllc 
'Chaenactis arlemisii/olia (A. Gray) A. Gray 
'Chaenactis glabriuscula DC. var. 'glabriuscula Golden curls 
'Chaenactis glahriuscula DC. var. lanosa (DC.) H.M. Hall 
"Cltaenaclis parishii A . Gray Parish 's pi ncushion 
'Cliamomilla suaveloens' (Pursh) R\db. Pineapple weed 

'Chrysothamnus nauseusus (Palla.s) Bntton subsp. bernardinus (H.M. Hall) H.M. 
Hall & Clements Gravelly cla\ soils, Cuyamaca Peak; Rabbit brush 

Hirshberg & demons: Cuyamaca and Laguna Mountains llora 81 

'Clirysolhanmus nauseosus (Pdllixs) Britton subsp. cotisiniiUs (E. Greene) H.M. Hall 

& Clements 
^CicJioriuin intybits' L. Mediterranean chicorv' 

^Cirsium occidenlale (Nutl.) Jcpson var. cali/ornicum (A. Gray) Keil & C. Turner 
^Cirsiiun scariosum Nutt. Bird's nest thistle 
*Cnicus henediclus' L. Blessed thistle 
^Conyza canadensis (L.) Cronq. Horseweed 
^Coreopsis californica (Nutt.) H. Sharsm. var. californica 
'Ericatneria brachylepis {A. Gray) H.M. Hall 
'Ericofneria cuneata (A. Gray) McClatchie var. macrocephala Urb. 
^Ericatneria linearifoUa (DC.) Urb. & J. Wussow 
'Ericatneria parislui {E. Greene) H.M. Hall var. parisliii 
'Ericameria pini/olia (A. Gray) H.M. Hall 
'Erigeron divergens Torrey & A. Gray Diffuse daisy 
^Erigeron foliosus Nutt. \<xx.foliosus Heabane 
"Eriopliyllum confertiflonim (DC.) A. Gray var. conferlifloritm 
^Eriophyllum wallacei (A. Gray) A. Gray Wallace's woolly-daisy 
^Euthamia occidenlalis Nutt. Western goldenrod 
*Filago arizonica A. Gray Poser and Viejas mountains (RSA-POM) 
^Filago californica Nutl. California filago 
^Filago gallica' L. Narrow-leaf filago 
^'Geroea viscida (A. Gray) S.F. Blake Sticky geraea 
"Gnap/wlium cali/ornicum DC. Green everlasting 
^Gnaphalium canescens DC. subsp. beneolens (Davidson) Stebb. & Keil 
^Gnaphalium canescens DC. subsp. niicroceplialum (Nutt.) Stebb. & Keil 
'Gnap/ialium paluslre Nutt. Lxiwland cudweed 
"Gnaplialium stramineuin Kunth Cotton-batting plant 
^Grindelia hirsulula Hook. & Arn. var. hallii (Steyerm.) M.A. Lane 
"Gulierrezia microcepluila (DC.) A. Gray Small-headed matchweed 
'Gutierrezia sarothrae (Pursh) Britton & Rusby Broom matchweed 
'Hazardia sqiiarrosa (Hook. & Arn.) E. Greene var. grindelioides (DC.) W. Clark 

Saw-tcx")th goldenbush 
"Heleniinn puberulum DC. Sncezewecd 
^Heliantliella californica A. Gray \ar. nevadensis (E. Greene) Jepson Near summit of 

Volcan Mountain 
'Helianthns annuiis' L. Western sunllower 
^Helianthiis californicus DC. California sunllower 
^Helianlhiis gracilenlus A. Gray Slender sunflower 
'Hetnizoniafa.sciculala (DC.) Torrey & A. Gray Golden tarweed 
"Helerollieca grandiflora Nutt. Telegraph ueed 

'Helerollieca sessiliflora (Nutt.) Shinners subsp. echioides (Bcnth.) Semple 
^Hieracixtn dlbiflonun Hcxik. While haukwecd 
^Hieraciiini (irgniiini Nutt. Golden hau kuccd 
^Hidsea californica Torrey & A. Gray San Diego hulsea 
^Hyinenopappus Jilifolius Hook. \ar. liigens {E. Greene) Jepson 
^Hynienoltiri.x wrightii A. Gray Wright's hymenolhrix 
'Hypocliaeris glabra' L. Smtx^th cat's ear 
'Isocoina nienziesii (Hook. & Arn.) Ncsom var. menziesii 
'Lacliica serriola' L. PnckK lettuce 

^Lagophxlla raniosissinia Nutt. subsp. ratnosissiwa Common harclcal 
^Laslhenia californica Lindlcy Calil'ornia goldlields 

82 PHYTOLOGIA August 1996 volume 81(2):69- 102 

'Lasllienia glaberrinia A. DC. Corte Madera and Cuyamaca lakes 

'Layia glandiilosa (Hcxik.) Hook. & Am. While tidy-lips 

'Lav/a platyglossa Fischer & C. Me>er Meadow tidy-tips 

*'Lessingia filaginifoUa (Hook. & Arn.) M.A. Lane \ar. filaginijolia Mountain 

'Lessingia glaiuiulifera A. Gray var. gUmduUfera 
^Maclioeraiuf^era (isteroides (forrey) E. Greene var. laguiieiisis (Keck) B. Turner 

Laguna aster 
"Mac/iaeranllwra canescens (Pursh) A. Gray var. canescetis 
^Maclmeranthera juncea (E Greene) Shinners Poser Mountain 
^Madia elegans Lindley subsp. elegaiis Elegant madia 
^Madia exigua (Smith) A. Gray Pygmy madia 
'Madia gracilis (Smi ih) Keck Slender madia 
'Madia minima (A. Gray) Keck Small madia 
'Malacolhrix clevelandii k. Gray 
'Micropus californicus Fischer & C. Meyer 
'Microseris douglasii (DC.) Schultz-Bip. subsp. platycarpfia (A. Gray) Chambers 

Viejas Mountain 
'Osmadenia tenella Null. Rosinweed 

"Pedis papposa Harvey & A. Gray var. papposa Cinchweed 
'Pentacluiela aurea Nutt. Golden daisy 
"Plucfiea odorata (L.) Cass. Salt-marsh fleabane 
'Psilocarphus brevissimus Nutt. var. brevissimns Woolly marbles 
'Rafinesquia californica Nutt. California chicory 
'Senecioflaccidus Less. var. douglasii (DC.) B. Turner & T. Barkley Sand-wash 

^Solidago californica Nutt. California goldenrod 
^Solidago confinis Nutt. Marsh goldenrod 
'Sonchus asper' (L.) Hill subsp. asper Spiny sow thistle 
'Sonchus oleraceus L. Common sow thistle 
'Stebbinoseris lieterocarpa (Nutt.) Chambers Viejas Mountain 
'Stephanomeria exigua Nutt. subsp. deanei (J.F. Macbr.) Gottlieb 
'Stephanomeria virgata Benth. subsp. pleurocarpa (E. Greene) Gottlieb Virgate 

'Stylocline gnaphalioides Nutt. Everlasting nest-straw 
^Taraxacum officinale' Wigg. Common dandelion 
^Tetradymia canescens DC. Monument Peak; Cotton-thorn 
'Telradymia cotnosa A. Gray 
'Tragopogon dubiiis' Scop. Goat's beard 
"Uropappus lindteyi (DC.) Nutt. Siher puffs 
*'Wyelliia ovata Torrey & A. Gray Mule's ears 
'Xanlhium siruinarium L. Cocklebur 

BERBERIDACEAE/Barbcrry Family 
'Berberis aqiiifoliiini Pursh \ ar. dictyota (Jepson) Jepson 
^Berberis aquifoliuni Pursh var. rf/^e-z/^ (Lindley) H. Scoggan 
'Herberts pinnala L^gasca subsp. pinnaia Shiny-leaf barberry 

BETULACEAE/Alder Family 
'Alnus rhonibifolia Nutt. While alder 

Hirshberg & demons: Cuyamaca and Laguna Mountains flora 83 

BIGNONIACEAE/Bignonia Family 

'Chilopsis linearis (Cav.) Sweet subsp. arcuata (Fosb.) Henrickson Desert willow 


'A/nsinckia menziesii (Lehm.) Nelson & J.F. Macbr. var. intermedia (Fischer & C. 

Meyer) Ganders 
'Cryptantim affinis (A. Gray) E. Greene Common cryplantha 
^Cryptanl/ui barbigera (A. Gray) E. Greene 
'Cryptantlui /loccida (Lehm.) E. Greene 
*Cryptantha intermedia (A. Gray) E. Greene Nievitas 
'Cryplantha micranl/ia (Torrey) l.M. Johnston 
'Cryptantlia micromeres {\. Gray) E. Greene 
*Cryptantlm microstachys (A. Gray) E. Greene 
*Cryptanlfia muricata (Hook. & Am.) Nelson & J.F. Macbr. 
'Cryptantfia simuUins E. Greene 

'Harpagonella palmeri A. Gray Poser and Viejas mountains 
*'Helioiropium curassavicum L. Salt helit)trope 
'Pectocarya penicillata (Hook. & Am.) A. DC. 
'Pectocarya setosa A. Gray 
*Piagiobotfirys arizonicus (A. Gray) A. Gray 
*Plagiobothrys bracteatus (J. Howell) l.M. Johnston 
'Plagiobotfirys canescens Benth. 
'Plagiobotlirys collinus (Philbr.) l.M. Johnston var. fulvescens (l.M. Johnston) 

'Plagiobotfirys hispidulus (E. Greene) l.M. Johnston 
^Plagiobotlirys nothofiilvus (A. Gray) A. Gray 
'Plagiobotlirys tenellus (Nutt.) A. Gray 
'Plagiobotlirys trachycarpus (A. Gray) l.M. Johnston 


^Arabis glabra (L.) Benth. var. glabra' Tower mustard 

^Arabis holboellii Homem. \ ar. pinetoriun (Tidestrom) Rollins 

^Arabis perennans S. Watson Nevada rock-cress 

^Arabis pulchra M.E. Jones var. graclis M.E. Jones 

^Arabis pidchra M.E. Jones var. pidchra Beautiful rock-cress 

^Arabis sparsijlora Torrey & A. Gray v ar. californica Rollins 

'Alhysanus piisillus {Hook.) E. Greene 

^Barbarea orthoceras Ledeb. Erect-pod wmter-cress 

'Brassica rapa' L. Field mustard 

'Camelina microcarpa' Andrz. Cuyamaca RSP; False llax 

'Capsella bursa -pastoris' (L.) Medikus Shepherd's purse 

^Cardamine californica (Torrev^ & A. Gray) E. Greene var. californica Milkmaids 

'Caidanllius lieterophyllus (Nun.) Payson var. heieropliyllus [C. stenocarpus Payson] 

Poser Mountain 
'Caulanlhus sinuilans Payson Gamet Mountain 
^Descurainia incana (Fischer & C. Mevcr) Dom 

'Descurainia pinnala (Walter) Bnlton subsp. glabra (Wooton & Standley) Detl. 
'Desciirainia pinnala (Waller) Bntton subsp. lialicloriini (Ccx^kercll) Detl. 
'Descurainia pinnala (Waiter) Bnlton subsp. menziesii (DC.) Detl. 
"Descurainia sophia' (L.) Webb Fine-leaf tansy-mustard 
'Draba cinieifoliaToncy & A. Gra\ Dcscn whitlow 

84 PHYTOLOGIA August 1996 volume 81(2):69- 102 

'Draba verna L. Vernal whitlow 

^Erysimum capilatum (Douglas) E. Greene subsp. capilatum 

'Guillenia lasiophylla (Hook. & Am.) E. Greene 

*'Hirsch/eldia incana (L.) Lagr.-Fossat Short-pod mustard 

"Lepidium campestre (L.) R.Br. 

'Lepidiutn lasiocarpumT oney & A. Gray var. lasiocarpum 

'Lepidium nitidum Torrey & A. Gray var. nitidum 

"Lepidiufn perfoliatum' L. Laguna Mountains 

'lepidium virginicnm L. var. pubescetis (E. Greene) Thell. 

"Raphanus raplianislnitn L. Mount Laguna; Jointed charlock 

"Rorippa curvisiliqua (Hook.) Britton Prostrate yellow-cress 

^Rorippa nasturtium-aquaticiun (L.) Hayek White water-cress 

^Rorippa simtala (Torrey & A. Gray) A. Hitchc. 

'Rorippa lenermmi E. Greene Lyrate-leaf yellow-cress 

'Sisymbrium altissimum L. Tall rocket 

'Sisymbrium officinale L. Tumble mustard 

"Streplanlhus bertmrdinus (E. Greene) Pansh Laguna jewelflower 

'Streplanthus campestris S. Watson Southern jewelflower 

'Thlaspi arvense' L. Corte Madera Ranch; Penny-cress 

'Thysanocarpus curvipes Hook. Lacepod 

'Thysanocarpus lacinialus Torrey & A. Gray Fnngepod 

'Tropidocarpum gracile Hook. Descanso; Slender dobie-pod 

CACTACEAE/Cactus Family 

'Echinocereus engelmannii (Engelm.) Lemaire Engelmann hedgehog 

'Opuntia basilaris Engelm. & J. Bigelow var. hasilaris 

'Opunlia clilorolica (Engelm.) J. Bigelow Pancake pear 

'Opunlia parryi Engelm. Cane cholla 

'Opunlia phaeacanlfia Engelm. Engelmann prickly-pear 

'Opuntia x vaseyi (J. Coulter) Britton & Rose Vasey pear 

CALLlTRlCHACEAE/Water-starwort Family 

'Callitriclie heterophylla Pursh var. bolanderi {V{e:^c\m.) Fassett 

'Callitriclie marginala Torrey 

'Callitriclie venui L. 

CAMPANULACEAE/Bell-Hower Family 

'Downingia concolor E. Greene var. brevier McVaugh 

'Gilhopsis diffusa A. Gray subsp, Candida (Euan) N. Morin 

'Gilhopsis diffusa A. Gray aubsp.fdicaulis (Ewan) N. Morin 

'Heterocodon rariflorum Nult. 

^lj)belia cardinalis L. var. pseudospknidens McVaugh 

Hjobelia dunnii E. Greene var. serrala (A. Gray) McVaugh 

'Nemacladus longiftorus A. Gray \ar. kmgiflorus 

'Neinacladus pinuatifidus E. Greene 

'Nenuuiadus rainusissiiiius Nutt. 

'Triodanis biflora (Ruiz Lopez & Pa\'on) E. Greene 

CAPRI FOLIACEAE/Honeysucklc Family 

^Lonicera subspicata H(xik. & Am. \ ar. dcniidnla Rchdcr 

'Sambucus mexicana C. Presl Blue eldcrbcrrv 

Hirshberg & Clemons: Cuyamaca and Laguna Mountains flora 85 

'Symphoricarpos mollis Nutt. Spreading snowberry 

'Symphoricarpos rotundifolius A. Gray var. parishii (Rydb.) Dempster Parish's 


^Cerastium fonlanum Baumg. subsp. vulgare' (Hartman) Greuter & Burdet Mouse- 
ear chickwccd 

*Cerasliu/n glomeratiun Thuill. 

'Dianthus arnieria L. subsp. anneria Volcan Mountain 

'Loe/lingia squarrosa (Nutt.) var. sqiiarrosa Poser Mountain 

*Mmuarlia douglasii (Toney & A. Gray) Mattf. Sandwort 

^Minuartia pusilla (S. Watson) Mattf. Little sandwort 

"Moehringia inacrophylla (Hook.) Fenzl 

'Sagina decumhens (Elliot) Torrey & A. Gray subsp. occidentalis (S. Watson) G. 
Crow Pearlwort 

^Saponaria officinalis' L. Bouncing bet 

'Silene antirrhina L. Snapdragon catchfly 

'Silene anneria L. 

*Silene gallicd L. Windmill pink 

^Silene laciniata Cav. subsp. major C. Hitchc. & Maguire 

^Silene lemmonii S. Watson Lemmon's pink 

*Silene mullinervia S. Watson Poser Mountain 

^Silene verecunda S. Watson subsp. platyola (S. Watson) C. Hitchc. & Maguire 

*Slellaria media' (L.) Villars Common chickweed 

'Stellaria niiens Nutt. Shiny chickweed 

CELASTRACEAE/Staff-tree Family 

'Euonymiis occidentalis Torrey var. parishii (Trel.) Jepson 


"Ceratophylliim demersum L. Corte Madera Ranch; Homwort 

CHENOPODIACEAE/Goosefoot Family 

'Alriplex rosea' L. Redscale 

'Clienopodium album' L. White lamb's quarters 

'Chenopodium herlandieri Moq. 

'Chenopodium californicum (S. Watson) S. Watson 

*Chenopodium hiatis Standley 

^Clienopodium incognitiim W'ahl 

'Salsola Iragus' L. Russian thistle 

CISTACEAE/Rcx-k-rose Family 
^Helianlliemum scoparium Nutt. Sun-rose 

CONVOLVULACEAEyMoming-glorv Family 
'Calystegia longipes (S. Watson) Brummit Piulc morning-glory 
^Cahsleqia macrostegia (E. Greene) Brummit subsp. arida (E. Greene) Brummit 
^Calystegia orcidenlalis (A. Gray) Brummit subsp. fulcrala (A. Gra\) Brummit 
Sonora morning glorv' 

86 PHYTOLOGIA August 1996 volume 8 1(2): 69- 102 

CORNACEAEVDogwcxxi Family 

'Coniiis nuttallii A ud ubon Paci f i c dogwood 

'Cornus sericea L. subsp. occidentalis (Torrey & A. Gray) Fosberg 

CRASSULACEAE/Stonecrop Family 

'Crassula aquatica (L.) Schonl. 

'Crassula contmta (Ruiz Lopez & Pavon) A. Berger Pygmy stonecrop 

^'Dudleya abrcunsii Rose subsp. abranisii Abram's dudleya 

^Dudleya edulis fNutt.) Moran Ladyfingers 

^Dudleya lanceolata (Nutt.) Bntton & Rose 

^Dudleya pulverulenta (Nutt.) Bntton & Rose subsp. arizonica (Rose) Moran Laguna 

^'Dudleya pulverulenta (Nutt.) Britton & Rose subsp. pulverulenta Chalk dudleya 
^Dudleya saxosa (M.E. Jones) Britton & Rose subsp. aloides (Rose) Moran [Dudleya 

alainae Reiser] 


"Cucurbita foetidissitna Kunth Calabazilla 

^Marah macrocarpus (E. Greene) E. Greene var. macrocarpus Wild cucumber 

CUSCUTACEAE/Dodder Family 

*Cuscuta califonnca Hook. & Am. var. californica Witch's hair 

'Cuscula ituiecora Choisy var. indecora 

*Cuscuta subinclusa Durand & Hilg. Canyon dodder 

DATISCACEAE/Durango Root Family 

*'Datisca gloinerata (C. Presl) Baillon Durango root 

DIPSACACEAE/Teasel^ Family 
^Scabiosa atropurpurea' L. Pincushions 

ELATINACEAE/Waterwort Family 

^Elatine hrachyspernui A. Gray Corte Madera Ranch 

^Elatine californica A. Gray California vvaterwort 

ERICACEAE/Manzanita Family 

'Arctostapliylos glandulosa Eastw. subsp. adamsii (Munz) Munz 

'Arctostophylos glandulosa Eastw. subsp. zacaensis (Eastw.) P. Wells 

'Arctoslapliylos glauca Lindley Big-bcrr\' man/iinita 

'Arclostaphylos pringlei C. Parry subsp. drupacea (C. Parry) P. Wells Cuyamaca 

'Arctostaphylos pungens Kunlh Mexican manzanita 
^Pyrota picia Smith Wintcrgreen 
'Rliododendron occidenlale (Torrey & A. Gray) A. Gray 
^Xylococcus bicolor Nutt. Mission manzanita 

^Cluiinaes\ce alhonmrginala (Torrey & A. Gray) Small 
'Chainaesyce nuiculala' (L.) Small Spotted spurge 
'Chaiuaeswe niicroniera (Engelm.) W(X)ton & Slandlcy 

Hirshberg & demons: Cuyamaca and Laguna Mountains flora 87 

*Chamaesyce serpyllifolia (Pers.) Small subsp. hirluUi (S. Watson) Koutnik Pine 

^Crolon californicus Muell. Arg. California croton 
^Eremocarpus setiger (Hook.) Benth. Doveweed 
^Eupliorbia palmeri S. Watson Wood spurge 
'Eup/iorbia spathulata Lam. Cuyamaca RSP 


'Acacia greggii A. Gray Cat-claw acacia 

'A/tiorpfia fruticosa L. False indigo 

^'Astragalus douglasii (Torrey & A. Gray) A. Gray var. parisfiii (A. Gray) M.E. Jones 

'Astragalus gambelianus E. Sheldon Poser Mountain 

'Astragalus oocarpus A. Gray Descanso milkvetch 

"Astragalus palmeri A. Gray Palmer's locoweed 

'Cercis occidentalis Torrey Western redbud 

^'Hoita macrostachya (DC.) Rydb. Leather root 

^Hoita orbicularis (Lindley) Rydb. 

^Lathyrus latifolius' L. 

^Lalhyrus vestitus Nutt. var. alefeldii (T. White) Isley 

^'Lotus argophyllus (A. Gray) E. Greene var. argopliyllus 

'Lotus crassi/olius (Benlh.) E. Greene var. crassifolius 

^Lotus grandi/lorus (Benlh.) E. Greene var. grandiflorus 

'Ij)tus hattiatus E. Greene 

^Lolus fieerniannii (Durand & Hilg.) E. Greene var. heennannii 

'Lotus humistratus E. Greene 

'Lotus micranthus Benth. 

^Lotiis nevadensis (S. Watson) E. Greene var. nevadensis 

^Lotus oblongifolius (Benth.) E. Greene \ar. oblongifolius 

'Lotus purshiamis (Benth.) Clements & E.G. Clements var. purshianus Spanish 

'Lotus saluginosus E. Greene \ ar. brevivexillus Ottley 
'Lotus saluginosus E. Greene \ ar. saluginosus 
^Lotus scoparius (Nutt.) Ottley var. brevialatus Ottley 
^Lotus scopariiLs (Nutt.) Ottley var. scoparius Deerweed 
'Lotus strigosus (Nutt.) E. Greene Bishop lotus 
'Lotus wrangelianus Fischer & C. Meyer 
'Lupinus albifrons Benlh. \ ar. albifrons 
'Lupinus bicolor Lindley Dove lupine 
'Lupinus concinnus J. Agardh 
'Lupinus excubilus M.E. Jones var. auslroinonlanus (A. A. Heller) C.P. Smith Grape 

stxla lupine 
'Lupinus excubilus M.E. Jones var. hallii (Abrams) C.P. Smith 
^Lupinus forniosus E. Greene \ ar. fonnosus 
'Lupinus liirsulissimus Benlh. Stinging lupine 
^'Lupinus latifolius J. Agardh \ ar. parishii C.P. Smith 
'Lupinus longifolius (S. Watson) Abrams Mesa Grande 
'Lupinus microcarpus Sims \ar. densiflonis (Benth.) Jepson 
'Lupinus sparsijlorus Benth. 
'Lupinus truncalus Hcxik. & Am. Chaparral lupine 
'Medicago liipiilina' L. Black mcdick 
'Medicdgo polyniorplia' L. Tixilhcd bur-ck)\cr 

PHYTOLOGIA August 1996 volume 8 1(2):69- 102 

*'Me(iicago saliva' L. Allalla 

"Melilolus alba' Mcdikus While sweet-clover 

'Melilolus indica' (L.) All. Indian sweel-clover 

'Pickeringia moiuatut Null. var. tonienlosa (Abrams) I.M. Johnston Cortc Madera, 

Viejas Mountains 
^Ruperiui rigida (Pansh) Grimes Pansh's rupertia 
'Spartium junceum L. Spanish broom 
^Thermopsis macrophylla Hook. & Am. var. semola Jepson 
'Trifolium albopurpureiun Torrey & A. Gray Rancheria clover 
*Trifolium ciliolatimi Benth. Tree clover 
'Trifoliu/n depauperalum Desv. var. iruiicatum (E. Greene) Isley 
'Tri/olium gracilenttdiriToTTey & A. Gray var. gracilenlum 
^Trifolium microcep/ialu/n Pursh 
'Trifolium obtiisiflorum Hook. & Am. Creek clover 
^Trifolium praterise' L. Red clover 
"'Trifolium variegatum Nutt. White-tip clover 
*Trifolium willdenovii Sprengel Tom-cat clover 
^Trifolium wonnskioldii Lehm. Cow clover 
^Vicia aniericana Willd. var. americana Sierra vetch 
^Vicia fiassei S. Watson Slender vetch 
'Vicia ludoviciana Nutt. var. ludoviciatia 
'Vicia villosa Roth subsp. varia' (Host) Corbiere 

FAGACEAE/Oak Family 

'Querciis agrifolia Nee var. agrifolia California live oak 
'Quercus agrifolia Nee var. oxyadenia (Torrey) J. Howell 
'Quercus berberidifolia Liebm. [Q. dutnosa Nutt. misappl.] 
'Quercus chysolepis Liebm. Canyon oak 
'Quercus corneliiis-mulleri K. Nixon & K. Steele 
'Quercus engeUnannii E. Greene Mesa blue oak 
'Querciis kelloggii Newb. Califomia black oak 
'Quercus wislizeiiii A.DC. \ar.frutescens Engclm. 

GARRYACEAEySilk-tassel Family 
'Garrya flavescens S. Watson Silk-tassel 
'Garryafremontii Torrey 
'Garrya veakiiii Kellogg 

GENTlANACEAEyCcntian Family 
^Cenlaiiriuin exallatum (Gnscb.) Piper Centaury 
^Cenlaurium venusluin (A. Gray) Robinson Canchalagua 
^Euslonia e.xattaliini (L.) Don Desert gentian 
'Swertia parryi (Torrey) Kuntzc Parry's green gentian 

GERANlACEAEyCeranium Family 

'Frodiiini bolrys' (Ca\.) Bcrtol. Long-beak filaree 

''Erodiiiin ciciitariuin' (L.) L'Her. Red-stem lilarcc 

\jeraiiiiiin califoniiciini G. Jones & F. Jones Agua Duicc Ra\ ine 

^Geraniiiin caroiniiaiiiiin L. Carolma geranium 

Hirshh)crg & Clcmons: Cuyamaca and Laguna Mountains fiora 89 

'Ril)es canl/iariforme Wiggins Moreno currant 
'Ribes indecorum Eastw. Winter currant 
'Ribes nevadense Kellogg Sierra currant 
'Ribes querceloruin E. Greene Oak gooseberry 
"Ribes roezlii Regl. var. roezUi 

m'DROPHYLLACEAE/Waterleaf Family 

*Emmenantlie penduliflora Benth. Whispenng bells 

'Eriodictyon tricliocalyx k.k. Heller var. lanatmn (Brand) Jepson 

'Eucrypla chrysanlfietnifolia (Benth.) E. Greene var. chrysanthemifolia 

*Nernophila nienziesii Hook. & Am. var. integri/olia Pansh 

'Nemopfiila menziesii Hook. & Am. var. menziesii 

'Netnophila pedunculala Benth. Cuvamaca Lake 

'Phacelia brachyloba (Benth.) A. Gray 

^Phacelia ciciitaria E. Greene var. hispida (A. Gray) J. Howell 

'Phacelia curvipes S. Watson 

^Phacelia davidsonii A. Gray 

'P/iacelia distans Benth. Wild heliotrope 

^Pluicelia imbricata E. Greene subsp. imbricala 

^Phacelia imbricata E. Greene subsp. palula (Brand) Heckard 

'Pfiacelia parryi Torrey 

^Phacelia raniosbsima Lehm. var. lalifolia (Torrey) Cronq. 

'Turricula parryi (A. Gray) J.F. Macbr. Poodle-dog bush 

HYPERICACEAE/St. John's Wort Family 
^Hypericum anagalloides Cham. & Schldl. Tinker's penny 
^Hypericum fonnosiim Kunth var. scouleri (Hook.) J. Coulter 
^'Hypericum perforatum' L. Klamath \v eed 


'Acanthomintlia ilicifolia (A. Gray) A. Gray San Diego thommint 
'Lamium amplexicaule' L. Henbit 
^Marrubium vulgare' L. Horehound 
^'Mentha arvensis L. Tule mint 

^Mentlm x piperita' L. Corte Madera Ranch; Peppermint 
^Mentlui spicata L. var. spicala' Spearmint 
"Monardella hypoleuca A. Gray subsp. lanata ( Abrams) Munz 
'Monardella lanceolata A. Gray Mustang mint 
"Monardella linoides A. Gray subsp. liiioides 
'Moiiardella luacraulha A. Gray Scarlet monardella 
"Monardella nana A. Gray subsp. leplosiphon (Torrey) Abrams 
"Monardella nana A. Gray subsp. nana 
^Pycnanllienuim calijornicuni Torrey Mountai n mint 
"Salvia apiana Jepson White sage 
^Salvia clevelandii (A. Gra\) E. Greene Fragrant sage 
^Salvia coliunbariae Benth. Chia 

'Salvia sononiensis E. Greene Vicjas Mountain, Cuyamaca Peak 
Scutellaria bolanderi A. Gray subsp. auslroinoniana Epling 
^Scutellaria tuberosa Benth. Blue skullcap 
*'Stacli\s ajii^oides Bcnlh. \ ar. ri^ida Jepson &. Hoov cr 

90 PHYTOLOGIA August 1996 volume 81(2):69-10: 

'Trichostema ausironionlanum Harlan Lewis subsp. aiLstromonUimim 
^Trichosletmi kmceolaluin Benth. Vinegar uced 
Trichostetm parisliii Vascy Mountain blue-curls 

LAURACEAE/Laurel Family 

'Umbellularia calijornica (Hcx^k. & Am.) Nutt. California bay 

LIMNANTHACEAE/Meadovvfoam Family 

'Limnanlhes gracilis Howell subsp. parisliii (Jepson) Beauchamp 

LINACEAEyRax Family 

*Hesperolinon micrantlumi (A. Gray) Small Dwarf flax 

^Linurn lewisii Pursh var. lewisii Blue Hax 

LOASACEAE/Stick-leaf Family 
'Mentzelia affinis E. Greene Blazing star 
'Mentzelia montana (Davidson) Davidson 
'Mentzelia veakhiaim Kellogg 

LYTHRACEAE/Ux^estnfe Family 

^Ly thrum californiciim Torrey & A. Gray 

'Lyihrum liyssopifolium L. Grass poly 

MALVACEAEyMallow Family 

'Malacothamnus deiisiflonis (S. Watson) E. Greene Bushniallow 
'Malva parviflora' L. Cheeseweed 

*'Sidalcea malvaeflora (DC.) Benth. subsp. sparsifolia C. Hilchc. 
^Spfiaeralcea ambigua A. Gray var. rosacea (Munz & I.M. Johnston) Kearney Mason 
Valley; Rose desert mallow 


^Melia azedaracli L. Banner China berry 


^Neliunbo lulea' (Willd.) Pers. American water-lily 

NYCTAGlNACEAEyFour-o'clock Family 

'Boerliavia intermedia M.E. Jones Ontlamme 

^Mirabilis californica A. Gray Wishbt'jnc bush 

^Mirabilis miiltiflora (Torrey) A. Gray var. piibescens S. Watson 

OLEACEAeOlive Family 
'Fra.xiniis velutina Torrey Velvet ash 

ONAGRACEAE/Evening Primrose Family 

"Cainissonia bisloria (Torrey & A. Gray) Raven 

'Camissonia californica (Torrey & A. Gray) Ra\'cn False mustard 

'Camissonia confiisa Ra\ en 

^Cainissonia liirlclla (E. Greene) Raven 

^Camissonia ignola (Jepson) Ra\ en 

^Camissonia pallida (Abrams) Ra\cn subsp. p(dli(la 

Hirshbcrg & demons: Ciiyamaca and Laguna Mountains llora 91 

*Caimssoiii(i slrii> (Fischer & C. Meyer) Raven 

^Clarkia delicala (Abrams) Nelson & J.F. Macbr. Corte Madera 

^CUirkia purpurea (Curtis) Nelson & J.F. Macbr. subsp. quiulrivultiera (Douglas) 

Harlan Lewis & M. Lewis 
'Clarkia rliomhoidea Douglas Diamond clarkia 
'Clarkia similis Harlan Lewis & Ernst Canyon clarkia 
'Epilohiuin brach\carpum C. Prcsl Summer cottonweed 
"Epilobium canuin (E. Greene) Raven subsp. canum 
"Epilobium canum (E. Greene) Raven subsp. lalifoUum (Hcx)k.) Raven 
^t'pHohium ciliatiun Raf. subsp. cilialum 
'Hpilobium densijlorum (Lindley) P. Hcx'h & Raven 
'Epilobium minutuin Lehm. Gravelly clay soils, Cuyamaca Peak 
*Epilobiutn py^inaeuni (Speg.) P. Hoch & Raven 
\iaura sinuala Ser. Buttertly weed 
^Gayophylum diffusum Torrey & A. Gray subsp. parviflurum Harlan Lewis & J. 

'Gayophylutn humile A.L. Juss. Cuyamaca Peak, Cuyamaca Lake 
*Ga\ophylum oligospennuin Harlan Lewis & J. Szweykowski 
*'Ludwii>ia repens Forster East ol Ramona; Water pnmrose 
^Oenothera californica (S. Watson) S. Watson subsp. californica 
\)enotliera elaki Kunth subsp. hirsulissinia (S. Watson) W. Dictr. 
^Oenolhera elala Kunth subsp. hookeri (Torrey & A. Gray) W. Dietr. & W.L. Wagner 

Anahuac Schcxtl; Great yellow evening primrose 

OROBANCHACEAE/Brtx)m-rapc Family 
^Orobanche bulbosa G. Beck Chaparral brtxim-rapc 
^Orubanclw fasciculala Nutt. Pine br(X)m-rape 
^Orobanche parishii (Jepson) Heckard subsp. parishii 


^Paeonia californica Torrey & A. Gray Calilbrnia peony 


"Argcnione nuinita Durand & Hilgard Prickly poppy 

'Dendroinecon rii^ida Benth. Bush poppy 

^Dicenlra chrysanllui (HcK)k. & Arn.) Walp. Golden car-drops 

"Eschscholzia californica Cham. Caiiloinia p<ippy 

'Meconella californica Toncy (, location doubtlul) 

'Mcconella denliculala E. Greene Mesa Grande (UCR) 

^Tapaver californiciini A. Gray Po.scr Mountain; Fire ptippy 

'Plaly.\tenu)n californicus Benth. Cream-cups 

lionnieya Irichocalyx E^stw . Matilija poppy 

^Planlai^o lanceolata' L. Rib-grass 
'Tlanldiio major' L. Common plant^im 
I'lanlaiio paldi^onica Jacq. Indian plantain 

Pl.A rANACLAiySycanu.ic lainil\ 
'l'l(il(itiii\ rai cinosa Null. Wcslcm swaiiunc 

92 PHYTOLOGIA August 1996 volume 8 1(2):69- 102 


'Allophyllum gilioides (Bcnlh.) A.D. Grant & V. Grant subsp. gilioides Straggling 

false gilia 
'Allophylluni gilioides (Benth.) A.D. Grant & V. Grant subsp. violaceum (A. A. 

Heller) Day 
'Allophyllum glutinosum (Benth.) A.D. Grant & V. Grant 
'CoUomia grandiflora Lindiey Grand phlox 
^Eriastrum densifolium (Benth.) H. Mason subsp. austromonlamun (Craig) H. Mason 

Chaparral woolly-star 
^Eriastrum fiUfolium (Nutt.) Wooton & Standley 
'Eriaslrum sapphirimtm (Eastw.) H. Mason Sapphire woolly-star 
^Gilia angelensis V. Grant Grassland gilia 
*GUia australis (H. Mason & A.D. Grant) V. Grant & A.D. Grant 
'Gilia capilala Sims subsp. abrotanifolia (E. Greene) V. Grant 
'Gilia caruifolia Abrams Garnet Peak, Stonewall Peak 
'Gilia clivoriim (Jepson) V. Grant 
'Gilia diegeiisis (Munz) A.D. Grant & V. Grant 

'Gilia ochroleuca M.E. Jones subsp. exilis (A. Gray) A.D. Grant & V. Grant 
'Linanthiis cilialiis (Benth.) E. Greene Whisker-brush 
'Linanlhus dianlhiflorus (Benth.) E. Greene Ground pink 
'Litianlhus dichotomiis Benth. Cuyamaca Lake; Evening snow 
'Linanlhus floribundus {A. Gray) Milliken subsp. floribundus 

'Litmntlnis floribundiis (A. Gray) Milliken subsp. glaber R. Patterson Summer snow 
'Linanlhus lemmonii (A. Gray) E. Greene Julian 
'Linanlhus liniflorus (Benth.) E. Greene 
'Linanlhus orcullii (C. Parr\' & A. Gray) Jepson 
'Litujnlhus parvijlorus (Benth.) E. Greene Baby stars 
'Linanlhus pygmaeus (Brdnd) J. Howell subsp. conlinenlalis Ra\en 
'Navarrelia atraclyloides (Benth.) Hcx)k. & Am. Skunkweed 
'Navarrelia inierlexla (Benth.) Hook. 
'Navarrelia peninsularis E. Greene Gravelly clay soils; Cuyamaca Lake, Cuyamaca 

'Navarrelia lageliiia E. Greene Meadows; Cuyamaca Lake 
"Phlox auslrunwnlana Cov . Pnckly phlox 
'Phlox gracilis E. Greene Slender phlox 

POLYGALACEAE/Milkwort Family 

'Polygala cornuta Kellogg var. fishiae (C. Parry) Jepson Viejas Mountain 

POLY GONACEAE/Buckwhcat Family 

'Chorizdiilhe fiiiibridia Null. \ar. laciiiiala (Toncy) Jepson 

'Chorizaiilhe leplolheca Goodman Corte Madera Ranch 

'Chorizanllu; polxgonoides Torre) & A. Gray \ar. longispina (Goodman) Miiiv. 

Knot weed spmcllowcr 
'Cliorizanlhe pronnnbens Null. Poser Mountain 
'Chorizdiilhe slalicoides Bcnlh. Turkish rugging 
'Eriogonuin apicutaluin S. Walson 
'Eriogoiuun davidsonii E. Greene 
'^Eriogonuin elongalnni Bcnlh. \ ar. clongaluin 
'Eriogonian fas(i(i(l(itiini Qcn[h. \m. Jusciculnluin Coilc Madera Ranch. Rat top 


Hirshbcrg & Clcmons: Cuyamaca and Laguna Mountains flora 93 

'Eriogonitin fcLSciculalum Bcnlh. \m. foliolosum (Nutt.) Abrams 

"Eriogonum fasciculatum Bcnth. var. poli/olium (A. DC.) Torrey & A. Gray 

'Eriogonum foliosumS. Watson Leafy buckwheat 

'Eriogonum gracile Benth. \'ar. gracile Slender buckwheat 

'Eriogonum molestuin E. Greene 

^Eriogonum nudum Benth. var. pauciflorurn S. Watson 

'Eriogonum parishii S. Watson 

'Eriogonu/n pluinaiella Durand & Hilgard 

*Eriogonutn thurberi Torrey 

'Eriogonum umbellalum Torrey var. munzii Rev. Mount Laguna 

"Eriogonum wrightii Benth. var. membranaceutn Jepson 

'Oxyt/ieca trilobata k. Gray Three-lobed spineflower 

^'Polygonum amphibiuin L. var. var. emersum Michaux Smartweed 

^'Polygonum amphibium L. var. stipulaceiun Coleman 

'Polygonum arenaslrum' Boreau 

'Polygonum douglasii E. Greene subsp. douglasii 

'Polygonum douglasii E. Greene subsp. yo/iAis/OAH/(Munz) J. Hickman 

'Polygommi lapathifoliion L. Willow smartweed 

'Polygonum parryi E. Greene Gravelly clay soils; Cuyamaca Lake, Cuyamaca Peak; 

Parry knotweed 
'Polygonum ramosissimuni Michaux Bushy knotweed 
'Pterostegia drynuirioides Fischer & C. Meyer Hair-net 
^Rumex acetosella L. Red sorrel 
^Rumex conglomeralus' Murray Green dock 
^Rumex crispus L. Curly dock 

^Ru/nex salicifolius J.A. Weinm. var. denticulat us Torrey 
^Rumex salicifolius J. A. Weinm. var. salicifolius Willow dock 

PORTULACACEAE/Purslane Family 

'Calandrinia ciliata (Ruiz Lopez & Pavon) DC. Red maids 

'Calyptridium moiuindrum Nutt. Sandcress 

'Claytonia exigua Torrey & A. Gray subsp. exigua 

'Claytonia pan'iflora Hook, subsp. parviflora 

'Claytonia parviflora Hook, subsp. viridis (Davidson) J.M. Miller & Chambers 

'Claytonia perfoliata Willd. subsp. mexicana (Rydb.) J.M. Miller & Chambers 

'Claytonia perfoliata Willd. subsp. perfoliata Miner's lettuce 

*Lewisia brachycalyx A. Gray Southwestern bitter-root 

^Lewisia nevadensis {A. Gray) Robinson 

^Montia chamissoi (Sprengel) E. Greene Cuyamaca, Laguna meadows 

'Montia fontana L. Water montia 

'Montia linearis (Hook.) E. Greene 

'Porlitlaca oleracea' L. Common purslane 

PRIMULACEAE/Pnmrose Family 

'Anagallis arx'ensis' L. Scarlet pimpernel 

'Androsace elongala L. subsp. acuta (E. Greene) G. Robb. 

'Centiinculus minimus L. Chariu eed 

^Samolus parviflorus Raf. Water pimpernel 


^Anemone luberosa R\db. Poser Mountain, Sco\e Canvon 

94 PHYTOLOGIA August 1996 \olumc Sl(2):69-102 

^Aquilegia fornu)sa Fischer Columbine 

^Clenuilis lasianlha Null. Virgin's boucr 

'Clenialis lignstki/olia Null. Verba de chnalo 

^Clematis j-Hiuciflora Null. Rope\me 

^Delphinium cardinale Hcx)k. Scarlcl larkspur 

^Delphinium lu'sperium A. Gray subsp. cuyamacae (Abrams) Harlan Leu is & Epiing 

Cuyamaca larkspur 
^'Delphinium parishii A. Gray subsp. subglobosum (Wiggins) Harlan Lewis & Epiing 

Garnel Mounlain, Pinyon Flal 
^Delphinium parryi A. Gray subsp. parry i 
^Delphinium patens Benth. subsp. hepalicoideum Ewan 
'M\osurus minimus L. Mouse-tail 
^Ranunculus aquatilus L. var. capillaceus (Thuill.) DC. 
^'Ranunculus aquatilus L. \ ar. hispidulus E. Drew 
^Ranunculus californicus Benth. California buttercup 
^Ranunculus c\mbalaria Pursh var. saximontanus Fern. 
'Ranunculus testiculatis' Crantz. Cuyamaca RSP (RSA-POM) 
^Tlmlictruinfendleri A. Gray Mn.fendleri Meadow-rue 
^Tluilicirum fendleri A. Gra\' var. polycarpum Torrey 

RHAMNACEAE/Buckthorn Family 

'Ceanothus cuneatiis (Hcx^k.) Null. var. cuneatus 

'Ceanothus foliosus C. Parry \ar. foliosus Viejas, Cuyamaca 

'Ceanothus greggii A. Gray \ar. perplexans (Trel.) Jepson 

'Ceanothus leucodermis E. Greene Whitebark wild lilac 

^Ceanothus oliganthus Null. \ ar. oliganthus 

^Ceanolhus palmeri Trel. 

^Rtuimnus crocea Null. Spiny redberry 

'Rhamnus ilicifolia Kellogg Holly-leaf redberry 

'Rlmmnus lomentella Benlh. subsp. lomenlella 

'Ziziphus parr\'i Torrey \ ar. parryi Lotebush 

ROSACEAE/Rose Family 

'Adenosioma fasciculatum Hook. & Am. Chamise 

"Adenostonui sparsifolium Torrey Red shanks 

^Agritnonia gryposepala Wallr. Agrimony 

^A/nelanchier utahensis Kc«hne ScrMce-bcrry 

^Cercocarpus hetuloide.s Tovrex & A. Gray \ar. beluloides 

^Cercocarpus miiuilijlnrus Abrams Viejas Mountain 

^Fragaria vesca L. Wild straw bcrrv 

^HeteroDieles arbulijolia (Lindlcy) R(.x?mcr Toyon 

^Holodiscus discolor (Pursh) Maxim. Cuyamaca Peak, Monument Peak 

^Horkeiui clevelandii {E. Greene) R\db. 

^Horketid irum ata Rydb. Viejas Mounlain, Boulder Creek 

^Polenlilla glandulosa Lindley subsp. gtandulosa Cinqueloil 

^Potenlilla glandulosa Lindley subsp. rejle.xa (E. Greene) Keck 

^Polenlilla gracilis H(xik. \ dr. fasti ^i(na (Null.) S. Wal.son 

Totentilla norvegicd' L. Rough cinqueloil 

Truiiiis enuir'^inata (Hook.) Walp. Bitter cherr\ 

Tniniis Iremonlii S. Banner; Desert apiicoi 

'Pridiiis iliiijolid (Null.) Walp. subsp. dici/otia 

Hirshberg & demons: Cuyamaca and Laguna Mountains flora 95 

'Pruniis virginiana L. var. demissa (Nutt.) Torrey 

*Rosa californica Cham. & Schldl. California wild rose 

'Rosa woodsii Lindlcy var. ullranumlana (S. Watson) Jcpson 

*'Rubns glaucifoliiis Kellogg var. gander i (L. Bailey) Munz 

^Ruhus laciniatus Willd. Cut-leaf blackberry 

^Rubus parvijlorus Null. Thimbleberry 

*'Rubus ursinus Cham. & Schldl. California blackberry 

^Sanguisorba minor Scop, subsp. miiricata' Briq. 

"Sanguisorba occidenlalis Nutt. Cuyamaca Lake, Cuyamaca Peak 

RUBIACEAE/Madder Family 

"Galium andrewsii A. Gray subsp. andrewsii Moss bedstraw 

'Galium angustifolium Nutt. subsp. angustifolium 

'Galium aparine L. Common bedstraw 

^Galium porrigens Dempster var. porrigens Oval-leaf bedstraw 

SALICACEAE/Willow Family 

'Populus fremontii S. Watson subsp. frefnonlii Fremont Cottonwood 

'Salix exigua Nutt. Narrow-leaf willow 

^Salix gooddingii C. Ball Black willow 

^ Salix laevigata Bebb. Red willow 

^Salix lasiolepis Benth. Arroyo willow 

SAURURACEAE/Lizard-tail Family 

'Anemopsis californica (Nutt.) Hook. & Am. Verba mansa 

SAXIFRAGACEAE/Saxifrage Family 

^Boykinia rotundifolia C. Parry 

^Heuchera brevistaminea Wiggins Ml. Laguna alum-root 

^'Heucliera rubescens Torrey var. versicolor (E. Greene) M.G. Stewart Cuyamaca 

Peak, Harper's Creek, Scove Canyon 
^Jepsonia parry i (Torrey) Small Poser Mountain, Cedar Creek 
^Litfwphragma affine A. Gray Woodland star 
^Ulliophragnia glabrum Nutt. Seeps, meadows; Corle Madera Ranch, Cuyamaca 

Lake, Laguna Mountains 
^Saxifraga californica E. Greene California saxifrage 

SCROPHULARlACEAEySnapdragon Family 

'Antirrhinum coulterianum Benth. Coulter's snapdragon 

'Antirrhinum kelloggii E. Greene Poser Mountain 

'Antirrhinum nuttalUanum Benth. subsp. nuttallianum 

^Castilleja affinis H(X)k. & Am. subsp. affinis Paint-bmsh 

^Ca.stilleja applegateiVcm. subsp. martinii (Abrams) Chuang & Heckard 

^Ca.stilleja altenuata (A. Gray) Chuang & Heckard 

'Castilleja densiflora (Bcnih.) Chuang & Heckard subsp. gracilis (Benth.) Chuang & 

^Castilleja exseria (A. A. Heller) Chuang & Heckard subsp. exserla \Orllwcarpus 

purpurascens Bcnlh.) 
^Caslilleja foUolosa HtKik. & Arn. WoolK paint-brush 
^Castilleja miniata Hook, subsp. miniala Giant red paint-brush 

96 PHYTOLOGIA August 1996 volume 81(2):69-102 

*CastiUeja minor (A. Gray) A. Gray subsp. spiralis (Jepson) Chuang & Heckard 
California thread-torch 

''Castiileja subinclusa E. Greene subsp. subinclusa 

*Collinsia childii A. Gray Child's blue-eyed mary 

'Coilinsia concolor E. Greene Southern Chinese houses 

^Collirisia heterophylla Buist Chinese houses 

^Coilinsia parvi/hra Lindley SmaJlflower blue-eyed mary 

^Cordylanthus nevinii A. Gray Cuyamaca Peak, Wooded Hill 

^Cordylanthus rigidus (Benth.) Jepson subsp. seiigerus Chuang & Heckard Thread- 
leaf bird's beak 

'Keckiella antirrhinoides (Benth.) Straw var. antirrliinoides 

'Keckiella cordifolia (Benth.) Straw Heart-leaf bush penstemon 

'Keckiella tertiata (Torrey) Straw var. lermta 

'Li/nosella acaiilis Sesse & Mocino Mudwort 

"Linaria canadensis (L.) Dum.-Cours. Blue toad-flax 

"Linaria genistifolia (L.) Miller subsp. dalmatica' (L.) Maire & Petitm. Dalmation 

'Mimulus androsaceus E. Greene 3 miles east of Banner 

'Mimulids aurantiacus Curtis [M aridus (Abrams) A.L. Grant; M. longijlorus (Nutt.) 
A.L. Grant; M. puniceus (Nutt.) Steudel] 

*Mimulus brevipes Benth. Slope semaphore 

^Mimulus breweri (E. Greene) Cov. Cuyamaca RSP 

^Mimulus cardinalis Benth. Scarlet monkey flower 

^Mimulus clevelandii Brandegee 

'Mimulus congdonii Robinson Damp, sandy areas in chaparral; Descanso, Poser 

'Mimulus floribundus Lindley 

'Mimulus fremonlii {Ben\h.) A. Gray 

'Mimulus gultalus DC. Seep monkey llovver 

'Mimulus palmeri A. Gray 

'Mimulus pilosus (Benih.) S. Watson 

'Mimulus rubellus A. Gray Indian Canyon, Poser Mountain (RSA-POM) 

^Pedicularis semibarbata A. Gray 

"Penstemon centranthifoliiis (Benth.) Benth. Scarlet buglar 

^Penstemon clevelandii A. Gray var. clevelandii 

"Penstemon tieleropliyllus Lindley var. auslralis Munz & l.M. Johnston 

"Penstemon heterophyllus Lmdley var. helerophyllus 

^Penstemon labrosus (A. Gray) Hook./ 

^Penstemon rostriflorus Kellogg [P. bridgesii A. Gray misappi.J 

^'Penstemon spectabilis Thurber var. speclabilis Showy penstemon 

^Scrophularia californica Cham. & Schldl. subsp. floribunda (E. Greene) Shaw 
California bee plant 

'Triphysaria eriantlui (Benth.) Chuang & Heckard subsp. eriantlm 

^Verbascum tlmpsus' L. Wcxilly mullein 

'Veronica anagallis-aqualica' L. 

'Veronica arvensis' L. Corn speedwell 

'Veronica peregrina L. subsp. xalapensis (Kunlh) PenncU 


'Ailanlhus allissima' (Miller) Sumglc Tree of heaven 

Hirshbcrg & Clcmons; Cuyumaca and Laguna Mountains flora 97 

SOLANACEAE/Nighlshade Family • 

"Ddlimi wrigluii Regcl Thorn-apple 

'Nicoliana atlenuala Torrcy Co\otc tobacco 

'Nicoliana glauca Grahiim Tree tobacco 

'Nicoliana oblusi/olia Martens & Galcolti Desert tobacco 

'Nicoliana quadrivalvis Pursh 

"Petunia pa rvijlora A.L. Juss. Wild petunia 

"Physalis crassijolia Benth. Thick- leaf groundcherry 

*Ph\salLs pul>escens L. var. inlegrifolia' (Dunal) Waterf. 

"Solanuni aniericanian Miller White nightshade 

'Solanum parisliii A. A. Heller 

'Solanum lanbeUiferum Eschsch. Blue witch 

'Solanum xanti A. Gray 


"Femonlodendron californicum (Torrey) Gov. subsp. californicwn 

STYRACACEAE/Storax Family 

'Styrax officinalis L. var. redivivus (Torrey) H. Howard 

TAMARlCACEABTamansk Family 

'Taniarix rainosissiiua' Ledeb. Corte Madera Ranch; Tamarisk 


'Celtis reticulata Torrey Net-leal' hackbenry 

URTICACEAE/Nettle Family 

'Hesperocnide lenella Torrey Corte Madera Ranch; Rock nettle 

'Parietaria Iwspera B.D. Hinton var. californica B.D. Hinton Poser Mountain 

^Urtica dioica L. subsp. holosericea (Nutt.) Thorne 

'Urlica urens' L. Corte Madera Ranch; Dwarl' nettle 

*Plecirilis hrachyslemon Fischer & C. Meyer 

'Plectrilis ciliosa (E. Greene) Jepson subsp. insignis (Suksd.) D. More\ Southern 

VERBENACEAE/Vervain Family 

'Phyla nodiflora (L.) E. Greene \ ar. nodiflora' Corte Madera 

^Verbena lasiostachys Link. \ar. lasioslach\s Western \er\ain 

VIOLACEAE/ Violet Family 

^Viola douglasii Steudcl Douglas \ lolct 

^Viold lobala Bcnth. subsp. integrifolia (S. Watson) R.J. Little 

^Viola lobata Bcnth. subsp. lobala 

*'Viola pediinciilala Torrcy & A. Gray Johnny-jump-up 

^Viola purpurea Kellogg subsp. purpurea Mountain violet 

^Viola purpurea Kellogg subsp. querceloruin (M. Baker & J. Clau.scn) R.J. Lilllc 

Wiola sororia Willd. subsp. affinis (LcContc) R.J. Little 

98 PHYTOLOGIA August 1996 volume 81 (2):69- 102 

VISCACEAE;Mislictcx; Family 
'Arceuthobiurn cali/oniicuni Havvksw. & Wiens 
''Arceulliobium ccunpylopodum Engelm. Western dwari mistletoe 
"Phoradendron detisiun Trel. Dense mistletoe 
^Phoradendron macrophyllum (Engelm.) Cockerell 
^Phoradendron pauciflorum Torrey White-fir mistletoe 
^Phoradendron villosum (Nutt.) Nutt. Oak mistletoe 

VITACEAE/Grape Family 

Wilis girdiana Mnnson Desert grape 

^Tribulus terrestris L. Puncture vine 


'Tree, 6 to 25 meters tall 

' Shrub, 0.5 to 6 meters tall 

' Chamaephyte, vegetative buds not over 0.5 meters above the ground 

*■ Hemicryptophyte, vegetative buds at surface of soil 

* Geophyte, vegetative buds below the surface of soil 


' Liana, vine 

■" Parasite 

" Plants with life forms ranging from annual to perennial or shrub. These are included 

in the life form spectrum as "unclassified". 
- Cryptogam (no code) 
' Denotes non-native taxon 
Cuyamaca RSP = Cuyamaca Rancho State Park 


The present checklist represents the work of many people. Duffie demons and 
Michael Curto prepared plant lists lor Palomar, Hot Springs, and the Cuyamaca and 
Laguna mountains ba.sed on their own extensive field experience. These lists were 
subsequently matched by Duffie demons with specimens in the collection housed at 
the San Diego Natural History Museum. Extensive collecting, especially of grasses, 
and taxonomic work by Linda Alien and Michael Curto have added much new 
information to the present list. Dr. Hcnk van der Werff's many collections from Corte 
Madera Ranch, Pine Valley, ha\c added new, interesting information, and filled 
in many gaps. The preparation of a synoptic collection lor San Dicgo County and the 
annotation of many specimens in the collection by Dr. Geoffrey Levin, curator of 
botany at the San Dicgo Natural History Mu.scuni from 19S4 lo 1 W3, has also added 
much new information. 

Hirshbcrg & Clcmons: Cuyamaca and Laguna Mountains llora 99 

Judy Gibson, Joan Stewart, Tom Obcrbaucr, Dr. Geoffrey Levin, and Steve Boyd 
provided many helpful commenLs on this manuscnpt. 


Allen, Linda S. 1994. Morphometries of Motiardella subgenus Macraiuluie 

(Lamiaceae). M.Sc. thesis, unpublished MS. California Polytechnic State 

University, San Luis Obispo, California. 72 pp. 
Beauchamp, R. Mitchel. 1972. Floral Diversity of San Diego County, California. 

M.Sc. thesis, unpublished MS. California State University, San Diego, 

California. 73 pp. 
Beauchamp, R. Mitchel. 1986. A Flora of San Diego County, California. 

Sweetwater River Press, National City, California. 241 pp. 
Chamlee, Helen. 1972. Ferns of San Diego County. San Diego Society of Natural 

History, San Diego, California. 72 pp. 
Clemons, Duffie & Michael Curto. 1986. Plants of montane San Diego County. 

Unpublished MS. 38 pp. 
Crow, Elisabeth D. 1931. A Study in Distnbution of Southern California Plants. 

M.Sc. thesis, unpublished MS. Claremont Colleges, Claremont, California. 107 

Curto, Michael. 1995. Grasses of montane San Diego County above 1,0CX) meters. 

Unpublished MS. 4 pp. 
Cuyamaca Rancho State Park. 1993. Wildllowers of Cuyamaca Rancho State Park. 

Color brochure with photographs by Suzanne Bell. 
Cuyamaca Rancho Slate Park. 1994. Shrubs of Cuyamaca Rancho State Park. Color 

brochure with photographs by Suzanne Bell. 
Hickman, J.C, ed. 1993. The Jepson Manual: Higlier Plants of California. 

University of California Press, Berkeley, California. 1400 pp. 
Higgins, Ethel B. 1949. Annotated distnbutional list of the ferns and flowenng 

plants of San Diego County. Occ. Papers, San Diego Society of Natural History 

No. 8. 174 pp. 
Hirshbcrg, Jen, et al. 1984. Plants of Cuyamaca Rancho State Park. Unpublished 

MS. 1 1 pp. 
Lower, Steven. 1977. of Springs in the Analysis of the Groundwater System at 

Mount Laguna. M.Sc. thesis, unpublished MS. California State University, San 

Diego, California. 235 pp. 
Munz, P.A. 1974. A Flora of Southern California. University of California Press, 

Berkeley, California. 1086 pp. 
Obcrbaucr, Thomas A. 1993. Soils and plants of limited distnbution in the 

Peninsular Ranges. Fremontia 21(4):3-7. 
Raven, P.H. & D.I. Axclrod. 1978. Ongin and relationships of the California llora. 

Unnersity of California Publications in Botany 72:1-134. 
Simpson, Michael, el at. 1994. Checklisi of the Vascular Plants of San Diego 

County. Special Publication No. 1. SDSU Herbarium Press. San Diego, 

California. 81 pp. 

100 PHYTOLOGIA August 1996 volume 81(2):69- 102 

Skinner, Mark & Bruce Pavlik, eds. 1994. California Native FUuU Society's 
Inventory of Rare and Endan/^ered Vascular Plants of California. Special 
Publication No. 1 /5th edition. Sacramento, California. 338 pp. 

van der Werff, Henk. 1992. Plants of Corte Madera Ranch. Unpublished MS. 12 

Wiggins, Ira Loren. 1929. Rora of San Diego County, Calilomia. Ph.D. thesis, 
unpublished MS. Stanford University, Palo Alto, California. 888 pp. 


This list is divided into three parts based on plant distribution, climate, and 
geology. For an explanation of plant habitat, ranty, and endangcrment codes, please 
refer to the California Native Plant Society's Inventory of Rare atid Endangered 
Vascular Plants of California (Skinner & F^vlik 1994). Geographic codes follow 
Hickman (1993). 

A. Plants occurring in both the Laguna and Cuyamaca mountains and/or adjacent areas 
including Corte Madera and Volcan mountains at elevations of 1212 to 1970 meters 
(4,000 to 6,500 feet). 

Delphiniutn hesperium subsp. cuyatnacae List lB:2-2-3, CR/C2 LCFrs, Medws 

(mesic), typical subsp. from SnFrB 
Helianlliella californica var. nevadensis LCFrs, VFGrs, near summit of Volcan 

Mountain, disjunct from SN 
East/ienia glaberrinia Medws, Corte Madera and Cuyamaca lakes, disjunct from San 

Luis Obispo County 
Eimnanlhes gracilis subsp. parishii List lB:2-2-3, CE/C2 Medws (vemally mesic), 

VnPls, typical subsp. from OR 
Minuartia pusilla Chprl, LCFrs/sand or clay, disjunct from SnFrB 
Thermopsis californica var. seniola List lB:2-2-3 CmWId, LCFrs, VFGrs, typical 

subsp. from nSW 

B. Plants occurring in the Cuyamaca Mountains at elevations of 1212 to 1970 meters 
(4,000 to 6,500 feet). 

Aspidotis densa LCFrs, Medws/gabbro outcrops on Cuyamaca Peak, disjunct from 

SN, usually on serpentine 
Ceanothus foliosus \m. foliosus Chprl/gabbro, di.sjunct from SnFrB 
Danlhonia unispicata Medws/gabbro, disjunct from SN 
Downingia concolor var. hrevior List lB:3-3-3, CE/Cl Medws (mesic), VnPIs, 

typical subsp. from SnFrB 
Epilobiuin minutuni Chprl, Medu.s/gabbro, Cu\amaca Peak, disjunct from CW, SN 
Gayophyluin huniile Mcdus/gabbro, Cuyamaca Lake and Cuvamaca Peak, disjunct 

from SnBr 
Ij^wisia hrachycalyx List 4: l-l-l Mcdus (mcsic)/gabbro, disjunct liom SnBr 
l^wisia nevadensis Medws (mcsic)/gabbro, disjunct from SnBr 

Hirshberg & Clcmons: Cuyamaca and Laguna Mountains tlora 


Navarrelia tagetina Mcdws, Cuyamaca Lake, disjunct from ScV 

Moehringia nuicrophylla LCFrs/gabbro, disjunct from SCoRO, usually on serpentine 

Polygonum parryi Mcdws (mesic)/gabbro, Cuyamaca Lake, Cuyamaca Peak, 

disjunct from SN 
Rubus glaucifolius var. gaiideri List lB:3-l-3 LCFrs/gabbro, typical subsp. from 

Sanguisorba occidentalis Medws/gabbro, disjunct from nCCO, SNH 

C. Plants occurring in the foothills of the Cuvamaca Mountains at elevations of 910 to 
1212 meters (3,000 to 4,000 feet). 

Ceanothus foliosus \ar. foliosus Chprl/gabbro, disjunct from SnFrB 

Filago arizonica Chprl, VFGrs/damp clay, coastal -desert disjunct, from sc AZ, nw 

Harpagotiella palineri List 2:1-2-1 Chprl, CoScr, VFGrs/damp clay, coastal-desert 

disjunct, from AZ, nw Mexico 
Mimulus congdonii Chprl/damp sand, disjunct from SN and Ventura County 





























































PHYTOLOGIA August 1996 volume 81(2):69- 102 













































































































Phytologia (August 1996) 81(2): 103-107. 


B.L. Turner 
Department of Botany, University of Texas, Austin, Texas 78713 U.S.A. 


A new species, Perymeniiim basaseachicanum B.L. Turner, spec, 
nov., is descnbed. It is known by numerous recent collections and is 
seemingly confined to southwestern Chihuahua. It belongs to the herbaceous 
elements of Perytnenium and is closely related to P. oxycarphum. A key to the 
four herbaceous species of Perymenium is provided, along with a map 
showing the distnbution of P. basaseachicanum, and a photograph of the 

KEY WORDS: Asteraceae, Perymenium, Mexico, Chihuahua, systematics 

Perynienium is a relatively large genus, containing approximately 40 species, most 
of these shrubby elements largely confined to Mexico (Fay 1978). A few species of 
Perymenium are low jjerennial herbs, the stems slender and dying back after each 
growing season, arising anew from swollen tuber-like roots. With the addition of P. 
basaseachicanum, descnbed below, the genus is now known to have four such 
herbacet:)us species, one of these P. buplillialmoides DC. ha\ing five 
morphogeographical varieties (Turner 1988). The following key will distinguish these 
four species, and Figure 2 shows the distnbution of P. basaseachicanum, which does 
not o\erlap that of the other taxa, nor do the sc\eral species appear to inlergrade. 


Involucral bracts 2-seriaie, subequal, mostly (6-)7-12 mm long. P. Imphfhalmoides 

Involucral bracts 3-4 senate, imbncaie to subimbncate, mostly 4-6 mm long (2) 

2. Lea\es elliptic, conaceous, widest near the middle; Jalisco P. jaliscetuse 

2. Lxa\es ovale, herbaceous, widest near ihe base; Pacific slopes (Oaxaca tt) 
Sonora) and western Chihuahua (3) 


104 PHYTOLOGIA August 1996 volume 81(2): 103-107 

3. Leaves \silh blades ovate to ovate-elliptic, 1-2 times as long as wide, the petioles 4- 
20 mm long, widespread F. oxycarphiim 

3. Leaves with blades ovate-lanceolate, 2-3 times as long as wide, the petioles 0.5-2.0 
mm long; Chihuahua P. basaseachicanum 

PERYMENWM BASASEACHICANUM B.L. Turner, spec. nov. Figure 1. 
TYPE: MEXICO. Chihuahua; Cascada Basaseachic, South Overview to Rancho 
San Lorenzo Hotel, pine-oak woodlands in "sterile tuff and nch meadows, 2150 
m, 22 Jul 1986, P.S. Marlin. el al. s.n. (HOLOTYPE: TEX!; Isotype: ARIZ). 

Simile P. oxycarplio S.F. Blake sed foliis medicalium ovatis-lanceolatis 
(vice foliorum ovatorum aut ovatorum-ellipticorum), 2-3plo longioribus quam 
latioribus (vice l-2plo), et petiolis 0.5-2.0 mm longis (vice 4-20 mm longis). 

Perennied herbs 30-50 cm high. Stems erect, arising from underground, tuber-like 
roots, sparsely to moderately stngose. Midstem leaves mostly 3-5 cm long, 1.0-2.5 
cm wide; petioles 0.5-2.0 mm long; blades ovate lanceolate, 3-nervate from the base, 
2-3 times as long as wide, sparsely to moderately hispidulous below with rough hairs, 
the margins remotely dentate. Heads l-3(-5) in terminal open cymes, the ultimate 
peduncles slender, 3-10 cm long. Involucres 3-4 seriate, 4-6 mm high, 6-10 mm wide 
(pressed); bracts broadly ovate, coarsely short-hispid, scarcely herbaceous, except for 
the short obtuse apices. Receptacles convex, 2-3 mm across. Receptacular bracts 
folded-lanceolate, persistent, their apices acute, hispidulous. Ray florets, 8-11, 
pistillate, fertile; ligules yellow, 8-10 mm long, ca. 2 mm wide. Disk florets 30-40 
(estimated), yellow, 4-5 mm long, the narrow throat ca. as long as the limb, the lobes 
5, ca. 0.7 mm long. Achenes ca. 3 mm long; pappus of ca. 20 deciduous bristles 1-3 
mm long. 

Guachochic, NW of Norogachic at airport, ca. 1950 m, 23 Aug 1978, Bye 8819 
(NMSU,TEX); Mpio. Bocoyna, mesa W of Creel between Creel and Ri'o Oteros, ca. 
2360 m, 16 Oct 1977, Bye & Weber 8204 (LL); Mpio. Guadalupe y Calvo, NW side 
of Cerro Mohinora, ca. 12.5 m SW of Guadalupe v Calvo, 2470 m, Nesoin 6487 
(TEX); Cascada Basaseachic, 1 Oct 1986, Martin, el al. 42 (NMSU); 37.6 km SW of 
El Vergel on rtwd to Guadalupe y Calvo, ca. 2500 m, 27 Aug 1983, Nesom 4949 
(TEX); Norogachi, ca. 5000 ft., 18 Jul 1955, Penningion 678 (TEX); Cascada 
Basaseachic, ca. 1 km airline S of Cascada m "thin soil over rock at overlook," ca. 
2100 m, 3 Oct 1986, Spellenberg, el al. 9270 (TEX). 

This species is superficially similar to Peryineniu/n buphlluihnoides \ar. lenellum 
(A. Gray) McVaugh, but is readily distinguished from the latter by its small 3-4 
seriate, subimbncate to imbricate involucres. Fay ( 1978) examined only one sheet of 
the taxon {Pennitiglon 678 [TEX]) annotating this as "probably" P. b. var. lenellum. 1 
was content with this name until numerous additional collections showed that the 
involucral characters aie more like those of P. oxycarphinn S.F. Blake, a widespread 
herbaceous taxon of the Pacific coast slopes (Turner 1987), distributed from Oaxaca to 
Sonora, and readily distinguished from /'. basaseachUannin by its broadly o\ate 
blades borne upon uell-dcfincd petioles, as noted m the abo\c key. 


New Peryineniiim from Chihuahua 


>.'« .•A«<«i| 


v..;;«iMw- ; 




Figure 1. Scanning photograph of Peryineniiim basaseachicanum (holoupc). 


PHYTOLOGIA August 1996 volume 81(2): 103-107 

Figure 2. Dislnbuuon of Ferymeiuuin hmaseachicaniini (based upon coilecuons al 
LL, TEX). 

Turner: New Peryinenium from Chihuahua 107 

It should also be noted here that label data on the Pennington collection (cited 
above) states that the Tarahumar name for the taxon is NAWABORl (a genenc name 
used for plants having edible roots) and that the Indians ate these, either "cooked, or 


I am grateful to Gayle Turner for the Latin diagnosis, and to her and Justin 
Williams for reviewing the manuscnpt. 


Fay, J.J. 1978. Revision oi Perymenium .. .. Allertonia 1:235-296. 
Turner, B.L. 1988. New combinations in Perymenium with special reference to P. 
bupluhalmoides . . .. Phytologia 64:263-266. 

Phylologia (August 1996) 81(2): 108-1 14. 


Edward E. Terrell 

Department of Plant Biology, University of Maryland, College Park, Maryland 20742 



Recent taxonomic papers by B.L. Turner concerning Texan and Mexican 
Hedyotis and Houstonia are reviewed. Diagnostic characters for Hedyolis and 
Houstonia are discussed briefly. Two subspecies of Houstonia acerosa are 
recognized: subsp. acerosa and subsp. polyprenioides. A key and distribution 
map of H. acerosa are provided. Houstonia pahneri is retained without 
varieties pending further study. One of three new varieties of Hedyotis 
nigricam established by Turner (var. gypsophila) is accepted here as 
taxonomically valid. A new species, Hedyotis pooleana, created by Turner, is 
synonymized under f/. mullerae. 

KEY WORDS: Rubiaceae, Hedyotis, Houstonia, systematics, Mexico, Texas 

The following commentary discusses taxonomic data on Hedyotis and Houstonia 
published by B.L. Turner (1995a-d). His four papers in the July and August 1995 
issues of Phytologia, owing to a lag in publication for this journal, were published in 
spring 1996, only about two months prior to the publication on June 10 of my 
monograph on Houstonia (Terrell 1996). I am now taking the opportunity to comment 
on Turner's papers. 

First, I wish to comment on Turner's circumscription of Hedyotis and his 
placement of H. acerosa and H. palmeri in Hedyotis rather than Houstonia. In my 
monograph (Terrell 1996) I circumscribed Houstonia to include 20 North American 
species. Hedyotis includes a broad heterogeneous array of species mostly native to 
Asia, with North and South American species presently in Hedyotis considered more 
distantly related to and probably not congcncnc with the Asian species. My 
monograph provided illustrations of representative seed types demonstrating that 
Houstonia seeds arc always cralcriform, i.e., have a ventral cavity, concavity or 
depression. In the subgenus Houstonia, the seeds have a rounded ridgcless cavity; in 
the subgenus Chaniisnie, the concavities or depressions have a hilar ridge. In 
contrast, ihc vast majority o( Hedyotis species are non-craleriforin, thus lack a ventral 


Terrell: Taxonomic notes on HedyoUs and Houstonia 109 

cavity, concavity, or depression. Those very few species which have crateriform 
seeds have other features setting them distinctly apart from both genera, and their 
proper classirication is being studied. The seeds of OkienUuuiia arc also non- 
crateriform, but are quite different in being conic or trigonous and much smaller and 
more numerous. These three genera also differ from each other in basic chromosome 
numbers (see Terrell 1996) and are thus genetically isolated. 

Houstonia acerosa (A. Gray) Bentham & Hooker/ {Hedyotis acerosa A. Gray) 

It is sometimes difficult in classifying a species to know whether to emphasize the 
morphological extremes as subspecies or varieties, or whether to emphasize the 
existence of much morphological overlap as indicative of there being just one variable 
species without infraspecific entities. In the case of the vanable Houstonia acerosa 
(alias Hedyotis acerosa) previous authors have tended to recognize the existence of 
two varieties or subspecies. In 1996, impressed by the extensive intergradation, I 
circumscribed Houstonia acerosa as just one vanable species without infraspecific 
entities. The opposite extreme was followed by Turner (1995b), who recognized four 
varieties: Hedyotis acerosa var. acerosa, var. polypremoides (A. Gray) W.H. Lewis, 
var. potosina B.L. Turner, and var. tamaulipana B.L. Turner. 

After re-considering the variation in Houstonia acerosa, I now believe that despite 
considerable intergradation, the two morphological extremes (see key below) and their 
partial but definite geographic separation (Figure 1) are sufficient to support their 
recognition as subsp. acerosa (Texas to San Luis PotosO and subsp. polypremoides 
(New Mexico and western Texas to Chihuahua and Coahuila), as previously published 
in 1979, rather than as varieties. The differences between these two entities are on a 
level higher than the usual variety. In addition, the geographical separation in New 
Mexio) and adjacent lands is clearly marked, despite intergradation in western Texas 
and northern Mexico. 

Turner (1995b) stated that \'ar. acerosa and var. polypremoides differ (among 
other characters) in the presence or absence of down-cur\'ed stem hairs; however, I 
could not find these hairs and interpret the vesliture as usually densely puberulent with 
very short hairs. (It may be noted that in his key to Mexican species Turner [1995b] 
erroneously listed a '\ar. fasciculala" , which should have been listed as \ar. acerosa). 
The following synonymy is summarized from Terrell ( 1996); the types are cited in the 

Houstonia polypremoides A. Grav, Prcx:. Anier. Acad. Arts 21:379. 1886. Hedyotis 
polypremoides (A. Gray) Shinners, Field & Lab. 17:168. 1949. Hedyotis 
acerosa A. Gray \ar. polypremoides (A. Gray) W.H. Lewis, Ann. Missouri Bot. 
Gard. 55:31. 1968, nom. superfl. Houstonia acerosa (A. Gray) Bentham & 
HcK)ker/ subsp. polypremoides (A. Gray) Terrell, Brittonia 3 1 : 168. 1979. 
Houstonia polypremoides A. Gray var. higelovii Grccnman, Proc. Amer. Acad. 

Arts 32:291. 1897. Hedvolis acerosa A. Grav var. bigelovii (Grccnman) 

W.H. Lewis, Ann. Missouri Bol. Gard. 55:397. ' 1969. 


PHYTOLOGIA August 1996 volume 81(2): 108-1 14 

400 km 

H. acerosa 

subsp. acerosa 
subsp. polypremoides 

Figure 1. Distnbulion oi' Houslonia acerosa subsp. acerosa (circles) and subsp. 
polypremoides (squares). 

Terrell: Taxonomic notes on Hed\otis and Houslonia 1 1 

Key to subspecies oi Houslonia acerosa 

1. Stems densely leafy; intemodes usually 3-10 mm long; leaves mostly verticillate, 
fasciculate, numerous, often crowded and overlapping, acerose; (lowers and 

capsules sessile or on pedicels to 3 mm long, sometimes densely grouped 

subsp. acerosa 

1. Stems not densely leafy; intemodes usually 10-25 mm long; leaves opposite or 
some verticillate, not or some leaves fasciculate, not crowded or overlapping, 
linear or acerose; flowers and capsules on pedicels 3-22 mm long, the 
inflorescence often open and diffuse subsp. polypremoides 

The remaining two varieties of Houslonia acerosa are discussed as follows. 

Var. polosina B.L. Turner has low, pulvinate, mat-like plants with much- 
shortened intemodes and long corolla tubes (8-10 mm long), and occurs from 
southernmost Coahuila to San Luis Potosf. In 1996 I considered these low plants as 
part of a cline that northward has taller plants with coarser leaves and longer 
intemodes. In southem Texas there are collections that are somewhat transitional, 
with rather fine leaves and small stature (e.g., Lewis 5517, Pecos Co.; Orcutl 734, 
Terrell Co.; and Pahner 11057, Valverde Co.; these at US). In corolla tube length 
there appears so much overlap that this character is not diagnostic. For these reasons I 
cannot recognize this new vanety, and synonymize it under subsp. acerosa. The type 
specimen of Hedyotis acerosa, Wrighl 237 (see Terrell 1996), was collected in 
"Western Texas to EJ Paso, New Mexico" in 1849, and consists of plants like those 
cited above. 

Var. tainaulipana B.L. Turner differs from var. polypremoides in having corolla 
tubes mostly 3-4 mm long and calyx lobes 1.5-2.0 mm long, comp»ared to 4-7(-12) 
mm long and 2-5(-7) mm long. Turner ( 1995b) cited a type and two other collections 
from Tamaulipas. I was previously aware of these different-looking collections, had 
examined them in detail after borrowing them on loan twice from TEX, and cited and 
mentioned them (Terrell 1996:91), while including them in H. acerosa sens. lat. My 
study of them confirmed Turner's measurement for the corolla tube lengths, however, 
for calyx lobe lengths I found 1.0 to 2.6 mm. The three collections grew on habitats 
described as "short brush", "short brush on calcareous terraces", and "shale hill". The 
plants were generally rather short, wiry-stemmed, and had consistently spreading, 
dilTuse inflorescences. The calyx and corolla tube lengths are the only differences by 
which \ar. kvnaulipana differs from subsp. polypremoides. The stem and 
inflorescence characters of var. tamai(li[\via are similar to those of subsp. 
polypremoides; in fact, the Tamaulipan plants tended to have a closer resemblance to 
collections of subsp. polypremoides from the Da\ is Mountains, Jeff Da\ is County, 
Texas (e.g., Rollins & Chambers 2759 [US]). Although var. tamaulijxma cxvupics a 
disUnct area to the south of collections oi subsp. polypremoides, its morphological 
differences are minor and overlapping, and 1 prefer to synonymi/,e it under the \anable 
subsp. polypremoides. 

112 PHYTOLOGIA August 1996 volume 81(2): 108-114 

Houstonia palmeri A. Gray {Hedyolis palmeri [A. Gray] W.H. Lewis) 

This Mexican species occurs in Coahuila, Nuevo Le6n, and San Luis Potosi 
(Terrell 1996). Turner (1995c) described var. muzquizana from the Miizquiz area of 
northern Coahuila It differs from var. paltneri in having corolla tubes (6-)8-10 mm 
long and pedicels mostly 20-30 mm long, versus 4-5(-8) mm long and 5-20 mm long. 
In my treatment (1996) of Houstonia palnieri I recognized only one variable species, 
but commented (p. 94) that plants in the Miizquiz area have the longest corollas. 
Collections from the Miizquiz area that were formerly available to me did not seem 
sufficiently different from the many specimens from farther south to ment varietal 
status. Recent loans of var. muzquizana (two collections from TEX, one US) bear out 
Turner's measurements of corolla tube lengths, but show some overlapping in pedicel 
lengths in the two varieties. My presently limited sample leaves me reluctant to accept 
the existing morphological data as conclusive concerning var. muzquizana, pending 
further collecting of it. 

Hedyotis nigricans (Lamarck) Fosberg, Lloydia 4:287. 1941. 

The third species studied by Turner (1995a) belongs to Hedyotis, and was 
therefore not included in my Houstonia monograph. Hedyotis nigricans is a 
polymorphic species ranging from Ohio and Kentucky across the southeastern United 
States to Florida and the Bahamas, west to eastern Colorado and New Mexico, and 
south to central Mexico. Terrell (1986; under Houstonia) provided a partial 
synonymy, and discussed two varieties occurring in Rorida. Tenrell (1991) 
summarized names and other data for the North American species of Hedyotis. 

Hedyotis nigricans and several related species including H. mullerae Fosberg have 
non-crateriform, ellipsoid seeds with a centnc punctiform "hilum. Chromosome 
numbers in this group are knov^n only for H. nigricans (x=9, 10). Seed morphology 
in the group differs distinctly from those of other Hedyotideae. The chromosome 
number of Oldenlandia is generally x=9, but this genus is quite distinct from Hedyotis. 

Turner ( 1995a) established three new varieties m Hedyolis nigricans. Var. 
austrotexana differs in being papillose with "extended epidermal cells" on the calyces, 
stems, and leaves. Plants are simple-stemmed and non-bushy herbs 20-40 cm tall and 
occur in several counties in southern Texas. The var. papillacea from the panhandle 
and trans-Pccos Texas is similar to the preceding variety in the papillose vestiture, but 
the plants are sprawling low bushy herbs 5-15 cm. tall. 

A few specimens of these varieties were lent to me from TEX, and the BRIT 
herbarium was visited in March, 1996. The characters, espcciall) those concerning 
habit, separating these \arielies seem to me to be minor. In ihc course of studying 
Houstonia species 1 ha\e examined and collected many specimens of Hedyotis 
nigricans from Texas and elsewhere. Considering the great \anability and \Mde range 
of H. nigricans 1 do not recognize the distinctness o\ these two varieties, and refer 
them to synonymy. 

Terrell: TsLwnomic noies on Hedwiis and Housionia 113 

The third entity, var. g\psopliila B.L. Turner, occurs in northern Mexico. 1 have 
been familiar with these plants for some years, and would ha\e descnbed them as \ ar. 
""elliplifolid', had I been less of a procrastinator. The \anet\ has elliptical leases and 
plants are consistently decumbent and short. Turner descnbed the lea\es as 3-6 mm 
wide, 2.5-3.5 times longer than wide, and the calyx as being setose with thickened 
hairs. I recognize this as a disunct variety. I was unaware of a preference of these 
plants for gypseous soils, as numerous a^llections recorded by me do not mention 
such soils. 

Hedyotis pooleatia B.L. Turner 

Turner (1995d) descnbed this new species from the Dead Horse Mountains, 
Brewster Co., Texas. It is closely related to Hedyotis niullerae, a low, compact, 
subshrub of Coahuila. I previously examined a number of specimens of H. midlerae 
from ASU, ENCB, GH, NT, TEX, and US. I have found H. nudlerae to be rather 
vanable in leaf size and shape, leaJ' vestiture, and compactness. Unfortunately, the 
only characters selected by Turner to disUnguish H. pooleana have to do w ith the stem 
leaves, and these do not seem to be much different from some specimens of H. 
niullerae, consequently, I refer Hedyotis pooleana to the s\nonymy of the vanable H. 
niullerae. Turner staled that the Dead Horse Mountains are an extension of the Sierra 
del Carmen oi Coahuila, u here H. niullerae is known to occur. 


I thank B.L. Turner, Director of TEX herbarium, for a loan of selected specimens. 
I am also grateful to the curators of the other herbana cited in the text. I acknowledge 
helpful reviews by J. Hennckson and J.L. Reveal, and useful laxonomic data from 
Hennckson. This work was supported in p>art by FYoject F-12 of the Mar\iand 
Agncultural Expenment Station and Cooperative Extension SerMce. 


Terrell, E.E. 1986. Taxonomic and nomenclatural notes on Houstonia nigricans 

(Rubiaceac). Sida 11:471-481. 
. 1991. Overview and annotated list of North Amencan species of 

Hedyotis, Houstonia. Oldenlandia, and related genera. Phytologia 71:212-243. 
. 1996. Revision of Houstonia (Rubiaccac-Hcdvoiideae). S\st. Bot. 

Monog. 48. 1 18 pp. 
Turner, B.L. 1995a. Taxonomic overview of Hedyotis nigricans (Rubiaceae) and 

closely allied laxa. Phytologia 79: 12-21. 
. 1995b. Taxonomv of the Hedyotis acerosa (Rubiaceae; complex. 

Ph\lolo2ia 79:83-88. 

114 PHYTOLOGIA August 1996 volume Sl(2): 108-114 

1995c. Taxonomic study of Hed\otis palnieh (Rubiaceae). Phytologia 


1995d. Hedyolis pooleana (Rubiaceae), a new species rrom the Dead 

Horse Mountains, Trans-Pecos Texas. Phytologia 79:93-96. 

I'liyloloi;iu (Augiisl IWO) 81(2) 115-117 


B.L. Turner 
Dcparlnicnt of Botany, Uni\crsily of Texas, Austin, Texas 78713 U.S.A. 


Venioiiia boqiieroiia B.L. Turner, spec, iiov., from Cerro Boqueron, 
Chiapas, Mexico, is de.scribed and pictured. It is reportedly an arching or 
clambering shmb to 3 m high, occuning on steep slopes with Finns, Qnercus, 
and Oslrva at about 2,255 m. 

KEY WORDS: Asteiaceae, Vemonieae, Vernonia, Mexico, Chiapas, 

Vernonia (s.l.) is a large genus with about 60 species in Mexico (Turner, in prep.). 
The present novelty was delected amongst a large collection of comps a.ssembled by 
Dennis Bi"ccdlo\e from throughout Mexico, all of these deposited at CAS. 

VERNONIA BOQUERONA B.L. Turner, spec. nov. Figure 1. TYPE: 
MEXICO. Chiapas: "Sleep slope with Finns, Qnercus dud Osirya on ridge NE of 
Cerro Boqueron on road from El Rosano to Niqui\il," 2,255 m, 29 Nov. 1986, 
D.F. lireedlove 66139, with J. Sigg (HOLOTYPE: CAS!). 

Similis Vernoniae pol\plenrae S.F. Blake scd frulex arcualus aul .scandens 
usque ad 2.5 m alius (vice fruticis aul arbons, ngide ereclae, 3-15 m allae) est, 
foliis caulibusque adpressis-pubcsccnlibus (\ice dense hirsutorum), et capitulis 
cum 8-9 llosculis (vice 20-22). 

Arching shrub or "vine" to 2.5 m high. Larger leaves linear-lanceolate to narrowly 
lanceolalc-ciiiplic, mostly 7-12 cm long, 1.6-2.6 cm wide; petioles 3-10 mm long, 
gradually pa.ssing into the blade; blades pinnalcly nervate, the ncncs prominent 
beneath, glabrale abcne, miiuilely pubescent to subglabrous below , the margins eiiliie 
or nearly st), the apices acute. Heads arranged in ralher compact to broadly 
lenninal corymbose panicles, the branches appics.sed-pubesccnt, clearly .somewhat 
arcuate uiiii ihe heads arranged along one side, sessile and ebracleate. the distance 



PHYTOLOGIA Autzusl 1996 \c)lumc 81(2); 115-117 

»woflf» c* ttturw, mrtm 

Vv? **•■■». '♦ ■"«• 

fS3«r»t'.ian S»M « 

Figure 1. Vernoiiia hoqticroiia ty[x: specimen. 

Turner: New Venioiiia from Chiapas 1 17 

between heads mostly 1-4 cm. Involucres turbocampanulate, 7-8 mm high, the bracts 
3-4 seriate, graduate, the inner series apprcssed-pubescent with silky hairs. Corollas 
purple, ca. 7 mm long, the lobes ca. 3 mm long. Achencs 8-9 per head; body 
somewhat 4-sidcd, ca. 4 mm long, 0.7 mm wide, pubescent throughout with 
ascendmg hairs; pappus double, an inner senes of ca. 40 dirty-white persistent bristles 
ca. 6 mm long, an outer scries of ca. 30 scales ca. 1 mm high. 

ADDITIONAL SPECIMEN EXAMINED: same collectors, site and dale as the 
holotypc but collection number 6(57 ;S(CAS,TEX!). 

The hololype has a \'er\' large open capitulescence (ca. 30 cm wide and as high), 
while Dreedlove 66118 has a much more compact capitulescence (ca. 14 cm wide, and 
as high), othenvise the two collections are very similar. 

Vernonia boqiteroiia has the general capitulescence and heads of V. [xjlyplenra, but 
differs markedly from the latter in habit, pubescence and florets per involucre, as noted 
m the above diagnosis. The latter species occurs in the same general region as V. 
/;o(7//<;'ro//rt but at seemingly lower elexations in wetter sites (mostly 1,200-2,200 m in 
montane rainforests based upon data from ele\en sheets at LL, TEX). 


I am grateful to Ga\le Turner for the Latm diagnosis, and to her and Ted 
Dele\ or\'as for re\iewing the paper. 

Phylologia (August 1996) 81(2) 118-120. 


B.L. Turner 
Dcpailmcnt of Botany, University of Texas, Austin, Texas 78713 U.S.A. 


A new species, Ageratina bobjaiisenii B.L. Turner, is described and 
illustrated from Duiango, Mexico. It belongs to the subgenus Neogreenella 
and is known only from holotypic nialcnal, this collected in pme-oak forests in 
the vicinity of Cd. Durango, 

KEY WORDS: Asteraceae, Eupaloneae, Ageratina, Mexico, Durango, 

Identification of a large assemblage of Mexican Asteraceae assembled by D.E. 
Breedlo\e from throughout Mexico (ca. 500 sheets [CAS]) has revealed the following 
novel tv. 

AGERATINA BOBJANSENII B.L. Turner, spec. iiov. Figure. 1. TYPE: 
MEXICO. Durango: "Rocky bluffs near Puerto Buenas Aires SW ol' L^ Ciudad 
in forest of Finns, Quercns and Arl>iiiii\' 2680 in, 7 Dec 1991, D.E. Breedlove & 
C. Burns 72708 (HOLOTYPE: CAS!). 

Similis A. znnil(nuie B.L. Turner ,scd habens folia ovala eglandulataque 
(vice loliorum cordatorum glandulalorumque) et in\'olucra pan'iora ca. 5 mm 
alia (\ice 6.5-8.5 mm). 

Stifny erect simple-stemmed perennial herbs 50-(30 cm high, the midstem 
internodes much longer than the leaves. Midstems moderately pubescent with short 
glandular hairs 0.02-0.03 mm high. Lea\es opposite throughout, at midslem 
mostly 6-9 cm long, 3-4 cm wide; petioles 2.5-3.5 cm long; blades broadly o\ale to 
somewhat deltoid, 3-nervatc from the base, sparsely pubcrulcnl with eglandular hairs, 
mostly along the edges and \eins, the margins crenulodcntalc. Heads numerous, 
arranged in a terminal capitulcscence uilh strictly a.scending branches, the ultimate 
peduncles 5-10 mm long, densely glandular-pubescent like the stems. Involucres 
narrow 1\- campanulatc, ca. 5 mm high, 2-3 mm wide (pressed); bracts 2-scnaie, linear- 



New A;^cniUna Ik mi Duiango 



N! "iSilOS 

Figure 1. A'^crniiiui hoh/diist'itii. Iroiii h()l(H\()C. 


PHYTOLOGIA August 1996 volume 81(2): 1 18-120 

lanceolate, eximbricate. Receptacle plane, glabrous. Florets 14-16 per head (two 
counts); corollas 5-6 mm long, pinkish-lavender, glabrous throughout, the tube ca. 3 
mm long, gradually tapering into the throat. Achenes ca. 2.5 mm long, sparsely 
ciliate; pappus of ca. 30 uneven ciliate, rather fragile bristles 3-5 mm long, the cilia 
mostly longer than the axis of the bnstles. 

This species belongs to the large subgenus Neogreemlla (ca. 100 species in 
Mexico) and will key to or near the Chiapasan species, Ageraliiia ziiiiilaiia B.L. 
Turner, in the treatment o( Ageraliiia in Mexico by Turner ( 1997). It differs from that 
species in being strictly herbaceous (50-60 cm high, vs. suffruticose or shrubby and 
ca. 1 m high), having mostly o\ate, glandular-pubescent leaf blades (vs. cordate and 
eglandular), and smaller involucres (ca. 5 mm high vs. 6.5-8.5 mm). 

There already being an Ageraliiia breedlovei King & H. Rob, 1 take pleasure in 
naming this novelty for my esteemed colleague Bob (Robert) Jansen, well known 
synantherologist cuirently engaged in studies on the phyletic relationships of comps 
generally, mainly using DNA data, but who, early on, produced a fine monographic 
study of the difficult genera Spilaiilhes and Acinella (Jansen 1981, 1985). 


I am grateful to Gayle Turner for the Latin diagnosis, and to her and Justin 
Williams for reviewing the manuscnpt. 


Jansen, R.K. 1981. Systematics of Spilaiilhes (Composilae: Heliantheae). Svst. 

Dot. 6:231-257. 
. 1985. The systematics of AcnieUa (Astcraceae, Heliantheae). Syst. Bot. 

Monogr. 8:1-115. 
Turner, B.L. 1997. Ageraliiia, In: Comps of Mexico Vol. 1: Ph\toiogia Memoirs 


Phytologia (August IW6) 81(2) 121123 


Joseph E. Laferriere 

Herbario, CEAMISH, Universidad Autonoma del Estado de Morelos, 62210 
Cuemavaca, Morelos, MEXICO 


Prionosciadium saraviki is a previously undescnbed species of ihe 
family Apiaceae. It is related to Prionosciadium townsendii but different from 
that species in several respects, pnmanly its yellow flowers, smaller fruits, 
and dentate rather than crenate lea\ es. 

KEY WORDS: Apiaceae, Prionosciadium, Mexico, Chihuahua, syslematics 


Prionosciadium saraviki es nueva especie de la familia de las 
Apiaceas. Es emparentada a Fr/oH05r/W/w/« townsendii pero difiere por tencr 
(lores amanllas, frutas mas pequenas, y hojas dentadas en vez de crenadas. 

PALABRAS CLAVE: Apiaceas, Prionosciadium, Mexico, Chihuahua, 

Prionosciadium saraviki Lafernere is a pre\ lously undescnbed species of the carrot 
family, called "saravique" or "saraviqui" by the Mountain Pima Indians of the Sierra 
Madre of Chihuahua, Mexico. The plant grow s in moist, shaded locations in its nali\ e 
habitat, which is a pine/oak forest with sabino (Cupressus lusilanica Mill.) frequent 
along creekbanks. The plant is valued for its edible root, which is white with a thin, 
papery, copper-colored periderm. The Mountain Pima scrape off the skin and slick the 
remainder of the rcx)t directly into the fire to roa-st it. 

As with many other umbellilcrs, sara\ique is a semelparous perennial. The Inst 
four or li\e years of ils life, it prexJuces only one or two leaves, each ol which is up U) 
90 cm across. When conditions are right, il forms a llowenng stalk up to 3 m high, 
bcanng hundreds of tiny \ellow tlowers. During 1987, a \ear ol adequate rainfall, 
approximately 20-307^ of the total population was in bkxim. the following \ear, a 
region was subject to an extreme drought, and as a result the onl\ indiv idual blcxmimg 


122 PHYTOLOGIA August 1996 volume 81(2): 121-123 

was one located in a permanent spring. The entire plant (mcludmg the rcxH) possesses 
a very pleasant aromatic fragrance. Saravique is becommg rare in iti> native habitat due 
to overutilization and overgrazing. 

In previous publications (Laferridre 1992, 1994a,b; Lafem^re, et al. 1991), I 
referred to the plant as Prionosciadium towiLsendii Rose. Further examination, 
however, reveals saraviqui to be different from that species in several respects, 
primarily its yellow (lowers, smaller fruits, and dentate leaves. Prionosciadium 
townsendii has purple llowers and crenate leaves. The Nabogame collections are 
therefore described below as a new member of the Apiaceae. 

Prionosciadium saraviki Laferr., spec. nov. TYPUS: MEXICO. Chihuahua: 
Municipio Tem6sachi, 2 km N of Nabogame, in spring on grassy sunlit hillside, 

18 km NNW of Yepachi, 28° 30' W, 108° 35' N, 1900 m, in llower, 27 Jul 1988, 
Laferriere 1569 (HOLOTYPE: ARIZ; Isotypes: ANSM,MEXU,UC). 

E Priotiosciadio townsendii Rose floribus luteis, foliis dentatis et fructibus 
4-8 mm longis differt. 

Plant semelparous, perennial, to 150 cm tall when fertile, bearing a pleasant 
aroma. Roots tuberous, with a pure white interior and a thin, papery, copper-colored 
periderm. Stems slightly reddish, waxy, to 4 cm in diam at base. Basal leaves ol' 
vegetative specimens 1-2, pentoid to deltoid in outline, pinnate, winged between 
leaflets, to 90 cm long, to 80 cm wide; leaflets pinnately lobed, the lobes lanceolate, 
dentate; petiole and rachis round, hollow. Upper cauline leaves opposite or whorled, 
temate or simple, with short, broad sheaths. Inllorescence of several verticils of 
peduncles; peduncles slender, 1.5-6.0 cm long, subtended by a pair of leaves, 
scaberulous at the summit; involucre wanting, or of a single filiform bract 5-10 mm 
long; involucel of several filiform bractlels 5-8 mm long, longer than the Howers but 
shorter than the fruit; fertile rays 8-16, spreading or divancate, subequal, 1.5-4.0 cm 
long; fertile pedicels 2-5, 3-7 mm long; calyx teeth obst:)lete; llowers yellow; ovary 
glabrous; petals inrolled at anthesis; anthers medially attached. Fruit oblong, truncate 
at the apex, retuse at the base, 4-8 mm long, 3-5 mm across, the dorsal ribs 
prominent, the lateral brcxidly thin-winged, the wings narrower than the btxiy; oil 
tubes 2 or 3. Seeds neady terete. 

Paratypcs: 2 km N of Nabcigamc, same individual as ht)lolype, m truit, 20 Qjt 
1988, /^/fmVrf 2756 (ARIZ); Nabogame, vegetative leaves only, 11 Jul 1988, I8(K) 
m, Laferriere 1504 (ANSM,ARIZ,MEXU); Nabogame, in llower, 18(X) m, 28 July 
1987, iM/erriere 535 (CHAPA,TEX,UC). 


Laferriere, J.E. 1992. Cultural and environmental response to drought among the 

Mountain Pima. Ecol. F(x)d Nutr. 28( 1 -2): 1 -9. 
. 1994a. Vcgctiilion and iloia ol ihc Mounlam I'lnia \illajjc ol Nalxiganic, 

Chihuahua, Mexico. Phylologia 77: 102- 140 

Lal'erriere: New Prionosciadium I romChihuahud 123 

1994b. A dynamic non-iincar optimi/ntion analysis of Mountain Pima 

subsistence patterns. Hum. E^'ol. 23(1); 1-28. 

, C.W. Weber, & E.A. Kohlhcpp. 1991a. Use and nutritional composition of 

some traditional Mountam Pima plant foods. J. Ethnobiol. 1 1(1):93-1 14. 

Phylologia (August 1996) 81(2); 124- 141 


Maria Elena Siqueiros-Delgado 

Centro bdsico, Universidad Autdnoma de Aguascalientes, Avenida Universidad s/n, 
Aguascalientes. Ags., 20100, MEXICO 

Yolanda Herrera-Ameta 

CIIDIR Unidad Durango, Institute Politecnico Nacional, Apartado Postal 57, 

Durango, Dgo., 34000, MEXICO 

Becana de la COFAA 


The study of anatomical characters in the culm epinodal zone of 41 species 
and sixteen varieties of Bouteloua has been completed. Worthless taxonomic 
characters were found in this region, nevertheless, five anatomical patterns 
were characterized. Other regions of the culm were explored for a possible 
later study. 

KEY WORDS: Bouteloua, Poaceae, anatomy, systematics 


Se llevo a cabo un estudio de los caracteres anat6micos en la zona epinodal 
del tallo de 41 espsecies y diesiseis vanedades del genero Bouteloua. Se 
encontraron caracteres de escaso valor taxonomico en esla region del tallo, no 
obstante, se caractenzaron cinco palrones anatomicos difercnies. Se 
complement6 el estudio con observaciones en olras regiones del tallo, para un 
p^isible estudio posterior. 

PALABRAS CLAVE: Bouteloua. Ptxiceae, anatom'a, sislematica 


Siquciros-Dclgado & Herrcra-Amcla: Bo ule loua analomy 125 


Systematic studies in plants are becoming more important up to date. From their 
results we can get a knowledge closer to the reality about evolution of the plants and 
their phylogcnetic relationships, particularly m those species with any kind of 
importance. This is the case of the Douleloua species, which are some of the most 
important native grasses in North American grasslands because of their great forage 
quality, their resistance to heavy livestock, and wide distribution. Species of 
Bouleloua were once the dominant components in the Mexican grasslands at the 
beginning of this century. However, in spite of being considered the producers of 
high quality beef in free-ranging livestock, their biology and taxonomy are not well 
defined until now, and the confusion in species definition remains. Thjs genus has 
been studied totally or partially by: Gnffiths (1912); Hitchcock (1920, 1935, 1939); 
Swallen (1939); Gould (1968, 1969, 1975, 1979); Gould & Kapadia (1964), and Roy 
(1968). All those studies were morphological taxonomic studies more than systematic 
ones. Gould, el al. incorporated chromosomal information in addition lo 
morphological data, but no other kinds of features have been studied in this genus in 
order to get an exact inference related to the evolution of its taxa. Columbus (1996) 
studied in his Ph.D. research, some aspects of a systematic study in this genus, 
together with other related genera. The topic of his research was the anatomy of leaf 
sections and molecular DNA in Bouteloua and genera related. From these two features 
he reached a conclusion about the phytogeny of the tribe Boutelouineae in America. In 
other studies, Herrera and colleagues began research to study basic features such as 
anatomy in leaf epidermis and culm sections, fiavonoid profiles, and physiographic 
requirements for the development of the Mexican taxa. 

The present study is part of an ongoing project "A biosystematic study of the 
genus Bouteloua from Mexico." At the same time it pretends to contribute to the 
Columbus research of Bouleloua from Amenca. The particular objective of this 
project is to charactenze worthy anatomical features from culm sections in the ta.\a of 
Bouteloua, in order to add data to support the better understanding of its phylogenetic 

It is widely known that anatomical studies in leaf sections add to the general 
knowledge of the plants with gotxl results (Ellis 1976; Columbus 1996; Herrera & 
Grant 1994; Aiken, et al. 1984). However, there are few studies of culm anatomy, 
none \n Bouteloua. Authors like Sanchez (1983a, 1983b), Ebinger & Carlen (1975), 
Auquier & Somers (1967), and De Wet (1960) have generated anatomical data from 
culm sections in several grass genera other than Bouteloua. All these authors agree 
about the phylogenetic value of culm anatomy, in addition to other features at the 
supraspecific level. 


The culm sections studied here ucrc made in the proximal node area (cpinodai 
zone) in 41 species and sixteen Nancliesof Bouleloua from Amenca, all ol ihcm Irom 
fresh collections done b\ Columbus (1996) (all the taxa \Mth numbers in this work. 

126 PHYTOLOGIA August 19% volume 81(2): 124-141 

e.g., B. aUuTiosam [2418]) in Mexico and other counlnes in America. 

Complementary fresh matenal collected by Herrera and colleagues., as well as 

herbanum specimens of rare species were used. Three hundred permanent slides with 

four to six repetitions per species, were done at the Anatomy Laboratory of Rancho 

Santa Ana Botanic Garden, Cluremont, Calif. The sections were cut at exactly the 

basal portion of the central internode of well developed culms, then processed as 


Desilicincation: 30% Hydrochloric acid was used for 48 hrs to remove silica from 


Dehydration: was in several consecutive steps of 1 hr each, in ethanol at different 

concentrations (30%, 50%, 70%, 90%), then 100% + 1% safranine to dye the tissues 

and to facilitate the inclusion and sectioning, then 100% ethanol alone. 

Clearing: consecutive steps 1 hr each were done in the following solutions: 2:1 

ethanol: xylene, 1:2 ethanol: xylene, 100% xylene twice. 

Impregnation: 1:1 Xylene: paraffin oil, 2 hrs; Paraffin at 58° C, 6 hrs, twice. 

Inclusion: was done in a special device to include tissue in the position required to 

make slices. 

Paraffin blocks with the tissue included were placed in a softener ( 10% aerosol 
OT: glycerine: water 10:3:90) from Schmid & Turner (1977), dunng two days in order 
to facilitate slicing. Blocks then were placed on ice at least 1 hr, then sliced with a 

Spencer 820 microtome. Slices of 10 |.im thickness were cut, then dyed following the 
Sharman (1943) procedure. Finally, the analysis and descnption of the anatomical 
characters was done. 

To get a comparative picture of the culm in Bouieloua, other regions than the 
epinodal were observed; these included the nodal zone and the central pxirt of the 
internode. Fresh material from specimens collected in several parts of Mexico by the 
collaborators of this project were used to analyze the differences and to validate the 
characters from other culm levels. 


The analysis of the fixed slices and the fresh slices permitted observation of the 
anatomy of the epinodal zone, as well as the nodal and central part of the internode 
(Figure 1), where it is possible to find several kinds of cells, organized in different 
ways, forming defined patterns depending pnmarily on the maturity and level of the 
culm. The main types of cells found arc: 

epidermal cells (Figure 2. 1), asscx-ialed with silica cells, trichomes and stomata; 
parenchyma cells, thin walled large cells which can form a medullar parenchyma 
(Figure 2.2), a subepidermal nng in the ncxlal region, or radial rachymorph 
parenchyma cells (chlorenchyma) which are responsible for culm photosynthesis 
(Figure 2.3), always placed perpendicularly to the epidermal surface and associated 
with the Kranz cells; 

sclerenchyma cells (Figure 2.4), thick walled cells with a small lumen forming 
subepidermal rings and generally surrounded by the va.scular bundles, making them 
more conspicuous at the apex; 

Siqueiros-Delgado & Hcnrera-Amela: Bo uleloua anatomy 127 

intermediate cells (Figure 4.1), morphologically intermediate cells which share 
features from both parenchyma and sclcrenchyma cells, very abundant in the culm and 
difficult to define, they form subepidermal nngs in proximal parts of the node, or they 
can be mixed in the sclcrenchyma ring. Some authors mention collenchyma cells in 
the culm, they are probably referring to this type of cells (Sdnchez 1981); 
mestome cells (Figure 6.4), thick walled cells that completely surround the 
peripheric vascular bundles and the intenor ones, they can be variable in size, number, 
and shape, and occasionally they look like sclcrenchyma cells; 

vascular bundles, can be internal (IVB, Figure 3.1) or peripheral (PVB, Figure 
3.2), the former are in the intenor part of the culm and are primary vascular bundles 
(Figure 3.3), occasionally secondary vascular bundles (Figure 3.4), while the later 
(PVB) can be secondar>' or even tertiary vascular bundles (Figure 3.5). Generally in 
well differentiated culm zones, the PVB are tertiary and are associated with Kranz cells 
and photosynthetic parenchyma, while in the immature zones the PVB are secondary 
vascular bundles; 

Kranz cells (Figure 3.6), large conspicuous cells, thin walled, associated with 
tertiary vascular bundles, in the exposed mature zones of the intemodal sheath. These 
cells together with the radial rachymorph chlorenchyma form a system called Kranz 
syndrome (Sanchez 1979), this system (Figure 3.7) is associated with the C^ 
photosynthetic process. 

Anatomical description of the culm internode at several levels 

Epinodal zone. Analysis of epinodal zone slides showed a rounded to oval 
transection, with smooth to lightly undulating outline. This culm zone of Bouteloua 
resulted in a very variable region for anatomical characters, however, it was possible 
to observe five well defined patterns in this zone, which are variable if the level of 
section is moved within just a few millimeters. 

1. Subepidermal ring of intermediate cells without sclcrenchyma. 

The subepidermal ring of intermediate cells which bound the PVB, these generally are 
secondary' VB, in some species showing an incipient Kranz system, the IVB are in one 
or several levels and can be primarv or secondary VB. The number and size of the VB 
is vanable among the species. Species containing this pattern are: Bouteloua 
almiosana [2418], B arislidoides var. aristidoides [2133, 2217], B. disliclui [2376], 
B. gracilis [2223], B. media [2367, 2420], and B. rigidiseta [2231] (Figure 4). 

2. Subepidermal ring of intermediate cells bounding a sclcrenchyma 
ring. The subepidcnnal ring of intermediate cells show the internal margin undulated, 
with 3 to 4 cells of thickness in the widest part, and two in the narrower. This nng 
surrounds a sclcrenchyma nng of two cells thickness that joins the VB, the PVB are 

totally included, they generally are IT, and bound the external portion the IVB can be 
pnmary or secondary. An incipient Kranz system is present in some samples. 
Species with this pattern are: Bouteloua ahvnosana [2301], B. barbata var. barbata 
[2268], B. chondrosioides [2422], B. curtipeudula \ar. caespitosa [2213, 2326], B. 
distidui [2393], B. elata [2366], B. eriostadiya [2286], B. media [2367, 2387], B. 

128 PHYTOLOGIA August 1996 volume 81(2): 124-141 

chihuafmana [2289], B. rigidisela [223 1, 2233), B. simplex [2278], B. iriaena [2338, 
2357], and, B. witliatnsii [2419] (Figure 5). 

3. Subepidermal ring of sclerenchyma alternated with intermediate 
cells. The subepidermal nng of sclerenchyma is variable in thickness, it surrounds 
the PVB and borders the external margin of the IVB, alternating with irregular zones 
of intermediate cells in some slides, where the ring is interrupted by Kranz cells with 
associated radial parenchyma. Species presenting this pattern are: Bouleloua atmua 
[2433], B. aristidoides var. aristidoides [2359], B. barbata var. barbata [2229, 2268], 
B. disticha [2360], B. elala [358], B. eludem [2274], B. eriopoda [2127, 2214], fi. 
gracilis [2340], B. hirsuta [2342], B. parry i var. parryi [2351], B. rejlexa [2436], B. 
scorpioides [2200], B. simplex [2335], B. trifida [2126], and B. williamsii [2353] 
(Figure 6). 

4. Ring of intermediate cells of sclerenchyma only in the top of 
vascular bundles. The nng of intermediate cells totally surrounds the PVB 

(generally 11°) with 3 to 6 thicknesses of cells, and borders the IVB only in the top. 
The sclerenchyma cells surround the VB in the top. Species presenting this pattern 
are: Bouleloua atuxua [2434], B. liirsula var. hirsuta [2284], B. tnegapotmnica 
[28393], B. parryi var. genlryi [2369], B. repetis [2123], and B. vaneedenii [2542] 
(Figure 7). 

5. Culm with well defined Kranz system. The sclerenchyma ring of 
variable thickness, interrupted by a variable number of Kranz cells like a crown, radial 
parenchyma is always associated, IVB in one or many levels. Species presenting this 
pattern are: Bouleloua aristidoides \2s. aristidoides [2217], B. barbaiavar. rotfirockii 
[2215], B. curtipendula var. tenuis [2352], B. pedicellata [2408], B. numsa [2287], 
B. repetis [2350, 2396, 2425], and B. iriaena [2401] (Figure 2). 

Observations done at nodal and intemodal zones from fresh sections of some 
Bouleloua species resulted as follows: 

Nodal zone. The nodal culm region presents a circular and smooth iranseclion 
with a subepidermal variable thickness parenchyma ring surrounding a thick 
sclerenchyma ring. The sclerenchyma ring links the VB's which are very close to each 
other. An unclear definition of the parenchyma nng and the VB's is found \n the initial 
node zone; sheath and culm are not well differentiated in this particular zone, appearing 
like a unit (Figure 8). This pattern is present in complete sequences of Bouleloua 
brevisela [2279] and B. rigidisela var. rigidisela [2231]. A dilferenl pattern in the 
same section level with external and internal sclerenchyma nngs was found in two 
other species: B. karwinskii [2208] and B. johiislonii [2285], where tvvo nngs of 
sclerenchyma are present, one external surrounding the parenchyma nng and another 
internal linking the undefined VB's. 

Examined species: Bouleloua aristidoides var. aristidoides [2279], B. barbata var. 
barbata [23\?,\,B. brevisela [2279], B. cliasei [2197], B. clwndrosioides [2184], B. 
curtipendula var. caespitosa [2177], B. eludens [2272], B. gracilis [2201], B. hirsuta 
[2216], B. karwinskii [2208], B. media [2367, 2387], B. peclinata [2823], B. 
purpurea [2343], B. radicosa [2318], B. ranu)sa [2137], B. rigidisela var. rigidisela 
[2231], B. scorpioides [23201. and B. uni^ora 12319]. 

Siqueiros-Delgado& Hcnrcrd-Ameta: Boute loua .inalomy 129 

Mid internodal zone. According to the observations from fresh or herbarium 
material slices, the anatomical pattern in this culm level (always devoid of sheath), 
seems to be constant in all the supenor half of the internodal zone, it matches with the 
Auquier & Somers ( 1967) description of a chlondoid type. There exist few variations 
depending on the species, mainly related to the number and position of the VB's, and 
the sclerenchyma nng thickness. The observations made in this zone were aimed at 
establishing differences among the anatomical features from the epinodal and the 
internodal zones, as well as to find important systematic characters. The descnplion of 
this pattern is shown as follows (Figure 3). Transections oval to round with smooth 
to undulate outline, epidermis with square or round cells, few tnchomes and stomata 
opening to the radial chlorenchyma. The subepidermal zone shows a continued 
sclerenchyma nng, interruptal by the Kranz system, this ring consists of 3 to 8 cells in 
thickness, surrounding the PVB, and bounding only the IVB top margin. The Kranz 
system with 4 to 8 crown shaped cells on the radial parenchyma, associated with the 
Iir VB; sclerenchyma girder associated with the PVB; VB's variable in number and 
position according to the sf>ecies, PVB's in this zone are mostly 111° with clearly 
associated Kranz system; I VB's can be distributed in one or several levels, they 
generally are 1°, sometimes 11°. The I VB's are always surrounded by their mestome 
sheath, which can have a single or double layer at the bottom. 

Examined species: Bouteloua alaniosana [Herrera & Siqueiros 1319] (CIIDIR), 
B. arislidoides var. aristidoides [Rosales sin] (HUAA), B. barbala var. barbala 
[Siqueiros sin] (HUAA), B. barbala var. sonorae [Herrera & Siqueiros 1309] 
(CIIDIR), B. clwndrosioides [Herrera 1402] (CIIDIR), B. curtipendula [Herrera 
1324] (CIIDIR), B. elata [Herrera 1465] (CIIDIR), B. parryi [Herrera & Siqueiros 
1315] (CIIDIR), B. pedicellala [Rzedowski 44714] (lEB), B. quiriegoensis [Herrera 
& Siqueiros 1320] (CIIDIR), B. radicosa [Herrera 1469] (CIIDIR), B. reflexa 
[Herrera & Siqueiros 1311] (CIIDIR), B. unijlora [Herrera & Siqueiros 1307] 
(CIIDIR), andfl. williatmii [Herrera 1300] (CIIDIR). 


The study of the Bouieloua culm anatomy resulted in two main problems: the 
enormous anatomical variation at different [evels, and the confusion and lower 
definition of some tissue cells. Metcalfe (1960) suggested that culm anatomy has a 
limited taxonomic value, because of the vanation at different levels, a theory which has 
been corroborated by the results of this work, in the epinodal purl oi the Bouieloua 

Anatomical features in the epinodal zone cannot be considered as key characters to 
add in the delimitation of Bouieloua species, neither in the determination of 
phylogenctic position; since it is a culm menstematic region, with different patterns at 
different levels. The five more stable patterns described in the results were based on 
the presence or absence and position of the sclerenchyma cells, as well as on the 
position of the ring of intermediate cells. The general observance for all the species is 

130 PHYTOLOGIA August 1996 volume 81(2): 124-141 

the presence of immature tissues (not completely differentiated) and a ling of 
intermediate cells bordenng the VB's, which is replaced by sclerenchyma cells far 
away from the node, Kranz cells are not present in this region. The VB's seem to be 
very mature in this zone, without variation from one region to another. It is difficult to 
establish a specific pattern inside the populations of one species, they commonly 
present intermediate schemes. The task of trying to assemble taxa related by 
anatomical features, and to compare them with phylogenetic groups previously 
established by Columbus (1996), did not succeed. Different taxa having the same 
pattern of culm anatomy do not represent close relationship (morphological, DNA 
content or leaf anatomy features). Unexpectedly, more than one culm anatomic pattern 
may be found inside populations of a single species, e.g., B. alamosana [2418] shows 
pattern # 1, while [2301] shows pattern # 2; B. disticha [2393] has pattern # 2, and 
[2360] has pattern # 3. This is probably due to millimetric differences in the level of 

Unexpectedly, the nodal zone was less variable, showing a specific anatomic 
pattern, which is easy to distinguish because it is very diffuse and the VB's are very 
close to each other, and formed a continuous ring of schlerenchyma which surrounds 
the entire VB. A subepidermal nng of parenchyma is often present, it rarely shows 
two sclerenchyma rings as observed in Bouleloua kanvinskii and B. johiistonii. This 
nodal pxittem changed towards upper levels, firstly substituting the typical parenchyma 
ring of this region for a ring of intermediate cells with a different arrangement (as 
commented above); then, in a higher level, this nng of intermediate cells was changed 
for sclerenchyma cells and the Kranz cells started showing up. The first cells appeared 
laterally placed, one on each side of the PVB, then two on each side, and so on until 
they completed a crown in the apex of the PVB. The complete crown was present al 
the uncovered zone of the sheath which corresponds to the central part of the 
intemode. Kranz cells were always associated with the radial parenchyma, which 
depending on the species can be conspicuous to inconspicuous. The presence of a 
mature Kranz system in this part of the culm, was probably due to the direct solar 
exposure, since it was devoid of the sheath protection at this portion, as Sanchez 
(1979, 1981) stated. At this culm level the resulting pattern remained constant in the 
reviewed populations. 

It is necessary to underline that the present study was based on the epinodal culm 
features, whose permanent slides were all done. The observations from higher levels 
of the intemode were done with fresh material, so that the quality was much lower, 
cind no key characters were determined, nevertheless it was possible to make general 
descriptions of the principal patterns. The number of repetitions per species was not 
enough to make generalizations, however, it was possible to get a precise idea of the 
culm anatomy in this particular region. It is obviously necessary to make a future 
study of the anatomical variation at the central part of the culm intemode. 

In spile of the gcxxl fixed slides obtained in this work, sometimes it was 
impossible to define the cells corresponding to the kind of tissue in observation, 
especially in the cpintxial zone, due to the intcmicdiatc cells mentioned and described 
above, which are morphologically intermediate between parenchyma and sclerenchyma 
cells, e.g., the cell wall thickness and the size. These cells have been catalogued by 
other authors as collcnchyma cells (Sanchez 19H4; Dc Wet 1960), or as tissue in 
transition or parenchyma in transition (Sanchez 1979, 1983a, and 1983b). Stevenson 
& Mertens ( 1980) slated ihal the collcnchyma and the parench>ina arc physically alike. 

Siqueiros-Delgado & Herrera-Arrieta: Bouteloua analomy 131 

since they both are bearing living and active protoplasts surrounded by pnmary walls, 
differing in the wall thickness and the length of the cells. The close relationship 
between these two tissues is very evident in the culms. Esau ( 1972) stated that in the 
adjacent region of collenchyma and parenchyma, there exist cells in transition, which 
share features from both cell types. Metcalfe (1960), mentions that there is frequently 
a transition between thin walled parenchymatous cells and thicker walled prosenchyma 
(sclerenchyma) cells. This transition can be gradual or abrupt and sometimes it is 
difficult to define the kind of cells under observation. Tissue in transition or 
intermediate cells are more common in the undifferentiated culm regions, where it is 
possible to find this tissue forming subepidermal rings, continuous or alternating with 
other tissues like sclerenchyma. When the alternating tissue is parenchyma, it is 
difficult to define its boundaries. The parenchyma cells from the culm anatomy are 
clearly defined as a radial parenchyma associated with the Kranz structure and to the 
medulla cells when present. 

During this exhaustive review of culm anatomical characters, searching for 
similarities and dissimilarities to define the taxonomic relationships, quite variable 
features were found at intra and interspecific level, e.g., the position of the ring of 
intermediate cells is not constant in the same individual at different culm levels; and 
also in different species. On the other hand, some features remain constant in the same 
species, but with minor variation between different species; that is the case of the VB's 
which present very low ranges of interspecific variation. 

Lastly, it is important to remark that the information generated by this study will be 
taken as part of the biosystematic study of the genus, and even though the initial 
hypothesis was not corroborated, the data generated are worthy, and contributes to the 
general knowledge of the genus Bouteloua. 


The main conclusion from this work is that the anatomic features from the epinodal 
culm zone in the taxa of Bouteloua, cannot be considered good systematic characters, 
because of the extreme variation presented. However, the central intemodal zone 
seems to be much more constant, unless additional studies suggest different 


This work was partially supported by CONABIO (B()6I) and CONACyT (3098- 
N). Special thanks arc given to Tra\ is Columbus for his kind advice and all the 
facilities to work with his freshly collected matenal, at his laboratory in Rancho Santa 
Ana Botanic Garden. 


PHYTOLOGIA August 1996 volume 81(2): 124-141 

Epinodal zone 
Nodal zone 

Mid internodal zone 
uncovered by the sheath 

Mid interodal zone 
covered by the sheath 

Epinodal zone 
Nodal zone 

Fig. 1. Levels of" culm sliced to observe; Nodal zone. Upinodal zone. Mid internodal zone 

Siqueiros-Delgado & Herrerd-Amcta: Bouieloiui anatomy 


Fig. 2. Bouteloua uniflora Vasey. Culm transversal section at the mid intemodal zone. 2.1) 
Epidermis. 2.2) Medullar parenchyma cells. 2.3) Radial parenchyma cells. 2.4) 
Sclerenchyma. 2.5) Vascular bundles. 2.6) Kranz system. 2.7) Internal vascular bundles. 


PHYTOLOGIA August 1996 volume 81(2): 124-141 

Fig. 3. Bouteloua pediceliata Swallen. Culm transversal section at the mid intemodal zone. 
3.1) Internal vascular bundles (IVB). 3.2) Peripheral vascular biuidles. 3.3) Primary 
vascular bundles. 3.4) Secondary vascular bundles. 3.5) Tertiary vascular bundles. 3.6) 
Kranz cells. 3.7) Kjanz system. 

Siqueiros-Delgado & Hcrrcrd-Amcla; Bouleloua anatomy 


Fig. 4. Bouteloua alamosana Vasey. Culm transversal section at the epinodal zone. 
Subepidermal ring of intermediate cells without sclerenchyma. 4.1) Intermediate cells. 4.2) 
Kranz cells. 4.3) Secondary peripheral vascular bundles. 4.4) Internal vascular bundles. 


HYTOLOGIA August 1996 volume 81(2): 124-141 

Fig 5 BoutelouaalamosanaVasey. Culm transversal section at the ep.nodalzone^ 
Subepidermal ring of mtermediate cells surrounding a ring of sclerenchyn.a. 5.1) Primary 
vascular bundles. 5.2) Secondar>' vascular bundles. 5.3) Kranz cells. 5.4) Ring of 
intermediate cells. 5.5) Ring of sclerenchyma. 

Siqueiros-Delgado & Herrerd-Amcta: Bouieloua anatomy 


Fig. 6. Bouteloua data Reeder & Reeder. Culm transversal section at the epinodal zone. 
Subepidermal ring of sclerenchyma alternating intermediate cells. 6.1) Intermediate cells. 
6.2) Ring of sclerenchyma 6.3) Peripheral vascular bundles. 6.4) Internal vascular bundles 
6.5) Medulla. 


PHYTOLOGIA August 1996 volume 81(2): 124-141 

Fig. 7. Bouteloua vaneedenii Pilger ex Urban. Culm transversal section at the epinodal 
zone. Ring of intermediate cells with sclerenchyma only at the apex of the vascular bundles. 
7.1) Intermediate cells. 7.2) Secondary peripheral vascular bundles. 7.3) Internal vascular 
bundles. 7.4) Sclerenchyma. 

Siqueiros-Delgado & Herrerd-Amcta: Bouteloua anatomy 




Fig. 8. Bouteloua radicosa (Foumier) Griffiths. Culm transversal section at the nodal zone. 
8.1) Ring of parenchyma. 8.2) Ring of sclerenchyma. 8.3) Secondary peripheral vascular 
bundles. 8.4) Internal vascular bundles. 

140 PHYTOLOGIA August 1996 volume 81(2): 124-141 


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Poaceae. Bull. Societe Royale Botanique de Belgique. Tome 100, p. 95. 
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Gould, F.W. 1969. Taxonomy of the Bouteloua repetis complex. Brittonia 21:261. 
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Station, Texas. 653 pp. 
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Gould, F.W. & Z.J. Kapadia. 1964. Biosystematic studies in the Bouteloua 

curtipendula complex. II. Taxonomy. Bnttonia 16: 182. 
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Sdnchez, E. 1981. Desarrollode la Eslructura Kranz en lallos dc Gramineae. Lilloa 

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Sanchez, E. 1983b. Dasyochloa Willdcnos\ ex Rydbcrg (Poaceae). Gcnero 
monoti'pico de Norteamenca. Lilloa 361; 13 1. 

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Phylologia (August 1996) 81(2) 142-143 


James L. Reveal 

Norton-Brown Herbarium, H. J. Patterson HaJl, University of Maryland, College 
Park, Maryland 20742-5815 U.S.A. 


The assertion that Acoraceae and Araceae (including Lemnaceae) form a 
group distinct from Arecaceae requires the establishment of a new subclass, 
Aridae. Its position within Liliopsida (monocots) is between Tnurididae and 
Liliidae. Arecidae is placed adjacent to Commelinidae. 

KEY WORDS: Liliopsida, Andae, nomenclature 

On-going phylogenetic studies on Liliopsida (monocots) necessitate some 
taxonomic realignments. The combined analysis by Chase et al. (1995) demonstrates 
that Arecidae {setisu Cronquist 1988) is polyphyletic. The new subclass includes 
Acoraceae and Araceae (including Lemnaceae). 

Aridae Reveal, subclass el slat. nov. based on the Latin descnption associated with a 
F.G. Banling {Ord. Nal. PL: 25, 65. Sep 1830, as Class Aropsida ["Aroideae"]) 

I accept the following subclasses in Liliopsida Scop., Fl. CarnioL: 228. 1760: 

k\\^mdX\&de'\-dk\\{., Syst. Fhylog. Magnolioph.: 461. 1967. (alismatoids) 
Tnurididae Takhl. ejr Reveal', Novon 2:235. 1992. (tnurids) 
Aridae Reveal, Phytologia 81: 142. 1997. (arids) 
]^\\\\&dtldk\\{.,Sysl. Phylog. MagnoUoph:. 473. 1967. (iilioids) 
kxcc\&deJdV.\\\.., Sysl. Phylog. Magnolioph.: 425. 1967. (arccids) 
Commelinidae Takhl., Syst. Phylog. Magnolioph.: 514. 1967. (commelinids) 
Zingibcridac Cronquist, Bnllonia 30:505. 1978. (/jngibers) 



New subclass in Liliopsida 



Work on this project has been supported by the University of Maryland 
Experiment Station and Cooperative Extension Service. This note was kindly 
reviewed by Drs. Bryan E. Dutton and Kancheepuram N. Gandhi. 


Chase, M.L., et al. 1995. "Monocot systematics: a combined analysis," pp. 685- 
709. In: P.J. Rudall, P.J. Cnbb, D.F. Cutler, & C.J. Humphnes (eds.). 
Monocotyledons: Systematics and Evolution. Kew, England. 

Cronquist, A.J. 1988. The Evolution and Classification of Flowering Plants. 2nd. 
edit. Bronx, New York. 





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